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53
UNIVERSITY OF CALIFORNIA PUBLICATIONS
ZOOLOGY
WILLIAM EMERSON RITTER
AND
CHARLES ATWOOD KOFOID
EDITORS
VOLUME XVII
WITH 29 PLATES
UNIVERSITY OF CALIFORNIA PRESS
BERKELEY
1916-1918
:
2
Ss ate yee
he 4
aj
:
‘e a=
10.
11.
CONTENTS
PAGES
. Diagnoses of Seven New Mammals from East-Central California, by
PeJiosepha Grinnelligand lira cype lim Loner eres ee eee ere 1-8
. A New Bat of the Genus Myotis from the High Sierra Nevada of
Calformiary bys Hilda Wood, (Grimm ell cence cece nee 9-10
. Spelerpes platycephalus, a New Alpine Salamander from the Yosemite
National Park, California, by Charles Lewis Camp ....................-..- 11-14
. A New Spermophile from the San Joaquin Valley, California, with
Notes on Ammospermophilus nelsoni nelsoni Merriam, by Walter
Tee ALOT estate tiscent fae se a a ee cae Rete ants ere cae a RNR eae Be sce oe 15-20
. Habits and Food of the Roadrunner in California, by Harold C.
Bryant) swath plates: as eee eae ee sensors are arene eee nese ence ee 21-58
. Deseription of Bufo canorus, a New Toad from the Yosemite
National Park, by Charles Lewis Camp ............0..2..2.2:.::essc1eseee-+ 59-62
. The Subspecies of Sceloporus occidentalis, with Description of a
New Form from the Sierra Nevada and Systematic Notes on
other California Lizards, by Charles Lewis Camp ....................-.-- 63-74
. Osteological Relationships of Three Species of Beavers, by F.
Harveyarloldentssswathiplatesy 2 ese eee ere areca ee eee 75-114
. Notes on the Systematic Status of the Toads and Frogs of Cali-
fornia, by Charles Lewis Camp :
A Distributional List of the Amphibians and Reptiles of Galetorntat
by Joseph Grinnell and Charles Lewis Camp
A Study of the Races of the White-fronted Goose (Anser albifrons)
Occurring in California, by H. 8. Swarth and Harold C. Bryant;
SWE TN eat UG 9 oo os aeons not scctce sa benesamuseresusecoseenecst ca cectem ed naeeeaseeeesee 209-222
. A Synopsis of the Bats of California, by Hilda Wood Grinnell; with
YO EEN SC STSp UCL ete ec ee ro ee ee ei eee 223-404
. The Pacifie Coast Jays of the Genus Aphelocoma, by H. 8. Swarth.... 405-422
. Six New Mammals from the Mohave Desert and Inyo Regions of
California, by Joseph Grinnell 423-430
. Notes on Some Bats from Alaska and British Columbia, by Hilda
Wood Grinnell 431-433
. Revision of the Rodent Genus Aplodontia, by Walter P. Taylor; with
DIBb eS PAO 29) eee ree 435-504
. The Subspecies of the Mountain Chickadee, by Joseph Grinnell...... 505-515
. Excavations of Burrows of the Rodent Aplodontia, with Observa-
tions on the Habits of the Animal, by Charles Lewis Camp ....... 517-536
UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol, 17, No. 1, pp. 1-8 ;
~ Vol. 17, No. 2, pp. 9-10 August 23, 1916
_ DIAGNOSES OF SEVEN NEW MAMMALS FROM
EAST-CENTRAL CALIFORNIA
BY
JOSEPH GRINNELL anp TRACY I, STORER
A NEW BAT OF THE GENUS MYOTIS FROM
THE HIGH SIERRA NEVADA OF
CALIFORNIA
BY
HILDA WOOD GRINNELL
UNIVERSITY OF CALIFORNIA PRESS
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Vol. 12. 1. A Study of a Collection of Geese of the Branta canadensis Group from
the Ban Joaquin Valley, California, by Harry 8. Swarth. Pp. 1-24,
plates 1-2, 8 text figs. November, 1913 on nner
2. Nocturnal Wanderings of the California Pocket Gopher, by Harold O.
Bryant. Pp. 25-29; 1 text fig. November, 1913 0 OE
3. The Reptiles of the San Jacinto Area of Southern California, by Sarah
Rogers Atsatt. Pp. 31-50. November, 1913 0c. | oD
4. An Account of the Mammals and Birds of the Lower Colorado Valley,
with Especial Reference to the Distributional Problems Presented,
by Joseph Grinnell. Pp. 61-294, plates 3-13, 9 text figs. March, 1914. 32.40
5. Aplodontia chryseola, a New Mountain Beaver from the Trinity Region
of Northern California, by Louise Kellogg. Pp. 295-296. ~
6. A Previously Undescribed Aplodontia from the Middle North Coast of
California, by Walter P. Taylor... Pp.. 297-300.
Nos. 56 and 6 in one cover. April, 1914 ni eee 05
7. A Second Species of the Mammalian Genus Miseviipoaons. from Call.
fornia, by Joseph Grinnell, Pp. 301-304. April, 1914 2.02. =O
8, Distribution of River Otters in California, with Description of a New
Subspecies; by Joseph Grinnell. Pp. 305-310, plate 14. October, 1914.05
9. Four New Pocket Gophers from California, by Joseph Grinnell. Pp.
911-816: «November, 19145223 2S oe Ss ne, * 05
10, Three New Races of Vespertilionid Bats from California, by Hilda é
Wood Grinnell. Pp. 817-321. December, 1914 2.2.28 OB
11. Eutamias sonomae, a New Chipmunk from the Inner Northern Coast
Belt of California, by Joseph Grinnell. Pp. 321-325, 1 text figure.
January; 1015 925-2 uct Se eS
ay,
2g AN TRANG RA ae
a PTC Ys ee
UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 1, pp. 1-8 August 23, 1916
DIAGNOSES OF SEVEN NEW MAMMALS FROM
EAST-CENTRAL CALIFORNIA
BY
JOSEPH GRINNELL anp TRACY I. STORER
(Contribution from the Museum of Vertebrate Zoology of the University of California)
Tn our attempts to determine the systematic status of the mammals
encountered in our natural history survey of the Yosemite section of
the Sierra Nevada several of the forms prove to have been heretofore
unprovided with names. It is the purpose of the present paper briefly
to characterize these new races, so that names will be available for
use in our distributional studies. Incidentally, related races in other
faunal areas have to a limited extent come in for remark.
The color names herein employed are taken from Ridgway’s Color
Standards and Color Nomenclature (1912). All measurements. except
of altitude, are in millimeters.
Scapanus latimanus campi, new subspecies
San Joaquin Mole
Type.—Male adult, skull and skin, in winter pelage; no. 21520,
Mus. Vert. Zool.; Snelling, 250 feet altitude, Merced County, Cali-
fornia; January 9, 1915; collected by Charles L. Camp; original no.
1746.
Diagnosis —Resembles Scapanus latimanus latimanus, but smaller,
pelage much paler and browner, feet and claws smaller, cranium
shorter, brain-case more inflated anteriorly, rostral region less taper-
ing, palatal region relatively wider, and supraoccipital ridge (posterior
to interparietal) higher. Resembles Scapanus latimanus occultus in
color, but decidedly larger in size, especially as regards feet and claws,
cranium heavier and much bulkier, brain-case deeper, and supra-
2 University of California Publications in Zoology [ Vou. 17
occipital ridge higher. Differs from Scapanus latimanus sericatus in
much paler and browner color, claws shorter and blunter, cranium
larger and broader, mastoid and interorbital breadths greater, and
brain-case more inflated.
Material—Three specimens from Snelling, Merced County, Cali-
fornia (nos. 22004, 22005, 21250, Mus. Vert. Zool.) ; one specimen
(injured skull and skeleton) from three miles north of Sanger, Fresno
County, California (no. 18854, Mus. Vert. Zool.). Both these localities
he in the Lower Sonoran Zone.
MEASUREMENTS (IN MILLIMETERS)
No. Sex Total length Tail Hind foot
21250 d 170 37 22
22004 Jb 170 i 36 22
22005 Jo juv. 155 35 20
Remarks.—The imperfect specimen from Sanger, by Jackson (1915,
p. 69) referred provisionally to occultus, seems to us to be better placed
with the form here newly characterized. It seems likely that campi
will be found to inhabit river-bottom lands of the San Joaquin Valley
generally, at least on the east side.
The form is named for Mr. Charles Lewis Camp, in recognition of
his ability as a field naturalist.
Martes caurina sierrae, new subspecies
Sierra Pine Marten
Type.—Male adult, skull and skin, in summer pelage; no. 22112,
Mus. Vert. Zool.; head of Lyell Canon, 9800 feet altitude, Yosemite
National Park, California; July 24, 1915; collected by Charles D.
Holliger; original no. 562.
Diagnosis.—Siunilar to Martes caurina caurina, but general colora-
tion paler both above and below, this paleness involving both overhair
and fur; sides of face decidedly paler; pale ochraceous-orange of chest
very extensive, spreading forward to throat and backward along
median line to belly; ‘‘feel’’ of pelage softer; cranium with rostrum
short as in caurina, but extremely narrow; whole cranium narrower,
and brain-case relatively higher; sagittal crest very weak; auditory
bullae even smaller than in cawrina.
Material—Nine skins with skulls, some also with body skeletons,
from various points in the Hudsonian Zone of the Yosemite National
Park, Mariposa and Tuolumne counties, California.
1916] Grinnell_Storer: Diagnoses of Seven New Mammals 3
MEASUREMENTS (IN MILLIMETERS)
Cranium
2 <2
Total Hind oi ae 2s 3es
No. Sex length Tail foot Ear 2 NS aa saa
22109 ref 600 180 80 35 69.9 47.6 36.0 29.3
22110 2 557 180 70 43 65.1 42.0 33.4 27.7
22111 3 640 215 86 BD. | (Ae ccceecty eee SS
22112 cf 609 190 79 35 71.0 46.6 35.8 29.1
22113 3 576 170 85 37 72.2 47.6 36.1 29.5
23040 ree 598 190 82 29 71.1 48.5 35.4 28.2
23041 djuv. 588 194 85 35 69.8 42.8 34.6 30.7
23042 2 551 177 76 33 65.8 43.8 33.7 27.0
23043 J 606 192 83 32 71.9 48.0 36.6 29.4
Remarks.—The Yosemite series of specimens, as regards both skins
and skulls, is notably uniform in characters, and is doubtless repre-
sentative of the race occupying the Boreal Zone along the whole Sierra
Nevada, north at least to Mount Shasta. The specimens referred to
caurina with some reservation by Miss Kellogg (1916, p. 356) should
now be referred to sierrae. They are not typical of that race, however,
showing departure in cranial characters towards cawrina proper.
Eutamias amoenus monoensis, new subspecies
Mono Chipmunk
Type.—Male adult, skin and skull, no. 23380, Mus. Vert. Zool.:
Warren Fork of Leevining Creek, 9200 feet altitude, Mono County,
California; September 25, 1915; collected by J. Grinnell; original no.
3709.
Diagnosis —Similar to topotypes of Hutamias amoenus amoenus,
in corresponding pelage (post-breeding), but general tone of coloration
paler, more grayish; top of head, central pair of light stripes on back.
and rump, with proportion of white to red greater; ochraceous-tawny
of sides paler; tips of dorsal tail hairs light buff rather than yellow-
ochre; light middle portion of ventral surface of tail ochraceous-buft
rather than ochraceous-tawny.
Material.—All from California: Twenty-one specimens from Mono
County: Walker Lake, 8000 feet; Silver Lake, 7200 feet; Gem Lake,
9036 feet; Mono Craters, 7500-8000 feet; Williams Butte, 7000 feet:
and Warren Fork of Leevining Creek, 9200-9400 feet; twenty-three
specimens from Placer County: Cisco, 6000 feet; Soda Springs, 6500
feet; twenty-six specimens from Nevada County: Independence Lake,
4 University of California Publications in Zoology [ Vou. 17
7000 feet. These localities all he in either the Canadian Zone or the
upper part of the Transition Zone.
Remarks.—The form here described is the southernmost and palest
representative of the amoenus group, occurring on the arid crest and
east wall of the central Sierra Nevada, where it is characteristic of the
Canadian Zone. Specimens from Independence Lake, Nevada County,
indicate intergradation toward true amoenus. Measurements of total
length, tail and hind foot, as tabulated by us (in MS), show no
significant differences.
Fifteen topotype specimens of Eutamias amoenus amoenus, taken
in the vicinity of Fort Klamath, Oregon, and loaned to us from the
collection of the United States Bureau of Biological Survey, through
Mr. E. W. Nelson, are in pelage comparable with our Mono series.
These proved invaluable for demonstrating the differences distinguish-
ing the race here newly named. é
Eutamias merriami mariposae, new subspecies
Mariposa Chipmunk
Type.—Female adult, skull and skin, in full winter pelage; no.
21855, Mus. Vert. Zool.; El Portal, 2000 feet altitude, Mariposa
County, California; November 24, 1914; collected by Walter P. Taylor;
original no. 7099.
Diagnosis—Sinilar to Eutamias merriami merriami in correspond-
ing pelage, but general tone of coloration grayer, less brownish; dorsal
pair of light stripes with ashy predominating; rump less brownish;
sides much grayer, with but a trace of the ochraceous-buft of merriami;
tail darker, the dorsal hairs banded and tipped with buffy white.
Material—Fifteen specimens from the Upper Sonoran and low
Transition zones of the west flank of the central Sierras of Cal-
fornia: Madera County: Raymond, 940 feet; Mariposa County ;: three
miles east of Coulterville, 3200 feet; El Portal, 2000 feet; Yosemite
Valley, 4000-5000 feet.
Remarks —The series at hand consists of specimens in either fresh
or worn winter pelage, with but two exceptions. One of these excep-
tions, a juvenal, as compared with merriama of the same age, is
decidedly paler and grayer, but not quite so gray as juvenals of
kernensis. The other exception, our only specimen showing summer
(post-breeding) pelage, is practically indistinguishable from merriami
in the same coat. It is thus quite different from the grayish kernensis,
though the tail is the same.
1916] Grinnell-Storer: Diagnoses of Seven New Mammals 5
Tn our attempt to establish the systematic status of the merriami
chipmunk of the Yosemite section, it was found that there exists an
additional previously undescribed race of the same group in the Kern
basin of the southern Sierras, and this is next described. Figure 1 is
a map showing the distribution of all the Californian races of
Eutamias merriami as illustrated by the material contained in the
- mariposae
- mM. pricei
-m. merriami
m. kernensis
Fig. 1. Map showing ranges of four races of Hutamias merriami
in central and southern California.
Museum of Vertebrate Zoology. This map, in conjunction with one
recently published by the senior author (Grinnell, 1915) will serve to
illustrate the ranges of all forms of the townsendi-merriami group
now known to oceur in California.
Eutamias merriami kernensis, new subspecies
Kern Basin Chipmunk
Type.—Male adult, skull and skin, in full summer (post-breeding )
pelage; no. 15022, Mus. Vert. Zool.; Fay Creek, 4100 feet altitude,
6 University of California Publications in Zoology [ Vou. 17
six miles north of Weldon, Kern County, California; July 13, 1911;
collected by H. A. Carr and J. Grinnell; original no. 266.
Diagnosis—Similar to topotypes of Eutamias merriami merriami
in corresponding pelage, but decidedly grayer in general tone of
coloration ; sides of head and hind-neck ashy gray rather than leaden
gray; dorsal light stripes relatively broad, more ashy in color; dorsal
dark stripes narrower, and less richly brown in shade; sides pale
tawny rather than reddish brown; dorsal tail hairs banded and tipped
with light buff rather than ochraceous-tawny ; ventral surface of tail,
centrally, ochraceous-tawny rather than cinnamon-rufous.
Material—F ifty-nine specimens from California as follows: Kern
County: Kern River at Bodfish, 2400 feet; Kern River, twelve miles
below Bodfish, 2000 feet; west slope Walker Pass, 4600 feet; Kern
River at Isabella, 2500 feet; Kiavah Mountain, 7000 feet; Fay Creek,
4100 feet; Tulare County: Taylor Meadow, 7000 feet; Trout Creek,
6000 feet; Jordan Hot Springs, 6700 feet. These localities all he in
the Upper Sonoran and Transition zones.
Remarks—AImost all of the specimens just mentioned are in full
summer pelage. Their general grayness is conspicuous as compared
with Eutamias merriami merriami, of which we have examined ade-
quate topotype material from the San Bernardino Mountains. Un-
fortunately winter specimens of kernensis are entirely lacking, so that
it is not possible to characterize the winter coat. A single July
specimen showing delayed molt suggests extreme grayness in a degree
comparable with that of the summer pelage.
Ochotona schisticeps muiri, new subspecies
Yosemite Cony
Type.—Male adult, skull and skin in winter pelage, no. 23480,
Mus. Vert. Zool. ; 9300 feet altitude near Ten Lakes, Yosemite National
Park, Tuolumne County, California; October 11, 1915; collected by
Walter P. Taylor; original no. 7720.
Diagnosis—Fairly intermediate between Ochotona schisticeps
schisticeps and Ochotona schisticeps albatus; general color more
brownish than in albatus, but not so dark as in schisticeps; head dis-
tinetly different from back, leaden gray as contrasted with lght
brown; under surface of body and upper surface of feet pervaded
with pale buff rather than either clay color or a creamy tint.
1916] Grinnell_Storer: Diagnoses of Seven New Mammals 1
Material—Fifty specimens from the Sierra Nevada in or near the
Yosemite National Park, in Mariposa, Tuolumne and Mono counties,
California. The various points of capture all lie within the Hudsonian
and Alpine-Arctie zones.
Remarks.—The extensive series of conies now available from the
Sierra Nevada points unmistakably to intergradation between schisti-
ceps and .albatus through the race here named muirt. Thus albatus
should henceforth be designated trinomially. Muiri is, if anything,
nearer to schisticeps of the Lake Tahoe region and northward than to
albatus of the Mount Whitney region.
The new form here characterized is named for a gifted Sierran
naturalist, the late John Muir.
Sylvilagus bachmani mariposae, new subspecies
Mariposa Brush Rabbit
Type—Male adult, skin and skull; no. 21867, Mus. Vert. Zool.;
adenostoma association, on McCauley Trail, at 4000 feet altitude, near
El Portal, Mariposa County, California; December 7, 1914; collected
by J. Grinnell; original no. 2972.
Diagnosis.—Similar to Sylvilagus bachmani bachmam, but general
tone of coloration decidedly grayer, more black on back, ears larger,
and cranium larger with longer and heavier rostral region; similar
to Sylvilagus bachmani cinerascens, but grayer, especially on sides
and ears, back with more black, general size greater, cranium with
longer and heavier rostral region, and auditory bullae conspicuously
smaller.
Material.—Six adult specimens from the Upper Sonoran Zone of
the western Sierran foothills in Mariposa County, California, 800 to
4000 feet altitude.
MEASUREMENTS (IN MILLIMETERS)
No. Sex Total length Tail Hind foot Ear from crown
21867 fot 326 30 73 80
21868 ie) 342 27 75 80
22928 2 340 32 al =
22929 fof 340 25 75 80
23619 ©) 280 25 75 75
23620 Jb 313 32 74 74
8 University of California Publications in Zoology [ Vou. 17
Remarks —The material at hand is probably representative of the
race of bachmani occupying the “‘lower slopes of the Sierras between
Shasta County and northern Tulare County,’’? which Nelson (1909,
6c
p. 249) refers to as ‘‘not typical’’ but in his estimation ‘‘nearest to
bachmani.’’
The several distinctive characters displayed by our speci-
mens seem to us to warrant fully the use of a separate name,
especially in view of the apparent complete separation of the ranges
of mariposae and bachmani. Tntergradation between these two forms
probably takes place through cinerascens, around the southern end of
the San Joaquin Valley. The latter race, however, is not at all inter-
mediate in characters, but is divergent from both, notably in the matter
of the enlarged auditory bullae.
LITERATURE CITED
GRINNELL, J.
1915. Eutamias sonomae, a new chipmunk from the interior northern coast
belt of California. Univ. Calif. Publ. Zool., 12, pp. 321-325, 1 fig.
in text.
JacKSON, H. H. T.
1915. A review of the American moles. U. S. Dept. Agric., Bureau Biol.
Surv., N. Amer. Fauna, 38, 100 pp., 6 pls., 27 figs. in text.
KELLOGG, LOUISE.
1916. Report upon mammals and birds found in portions of Trinity, Siskiyou
and Shasta counties, California, with description of a new Dipodomys.
Univ. Calif. Publ. Zool., 12, pp. 335-398, pls. 15-18.
NELSON, E. W.
1909. The rabbits of North America. U.S. Dept. Agric., Bureau Biol. Surv.,
N. Amer. Fauna, 29, 314 pp., 13 pls., 19 figs. in text.
Transmitted April 14, 1916.
UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 2, pp. 9-10 August 23, 1916
A NEW BAT OF THE GENUS MYOTIS FROM
THE HIGH SIERRA NEVADA OF
CALIFORNIA
BY
HILDA WOOD GRINNELL
(Contribution from the Museum of Vertebrate Zoology of the University of California)
Eight species of bats were obtained in the summer of 1915 by the
Museum collecting party working across the Sierra Nevada through
the Yosemite region. These are found to include an apparently un-
described race of the Yuma bat, which is herewith characterized. Of
particular note is the fact that this bat was ascertained to occur higher
zonally than any of the others, two specimens having been taken by
Dr. Walter P. Taylor at the upper edge of the Hudsonian Zone ( Vogel-
sang Lake, 10,350 feet altitude).
Myotis yumanensis altipetens, new subspecies
High Sierra Bat
Type.—M ale, adult ; no. 23034, Mus. Vert. Zool. ; 7500 feet altitude,
one mile east of Mereed Lake, Yosemite National Park, California;
August 19, 1915; collected by J. Grinnell; original no. 3437.
Diagnosis.—Largest race of Myotis ywmanensis occurring in Cali-
fornia (total length 88 to 93 millimeters; greatest length of cranium
14.2 to 15.0 millimeters) ; coloration nearest that of Myotis ywmanensis
sociabilis.
Material—F our specimens from the high Sierra Nevada, within the
Yosemite National Park: two from Mereed Lake, 7500 feet (Canadian
Zone), and two from Vogelsang Lake, 10,350 feet (Hudsonian Zone).
iQ) < University of California Publications in Zoology [ Vou. 17
MEASUREMENTS (IN MILLIMETERS)
Greatest
Total lengthof Weight
No. Sex length Tail Foot Ear Forearm Tibia cranium (grams)
23034 Jb 88 36 9 12 36:5." ess 15.0 7.8
23035 3 90 41 10 13 36.4 16.4 14.2 6.1
23036 2 91 37 10 14 36:0 5e eee 14.5 7.8
23526* J 93 40 11 15 36.0 IGG) 7 es 7.2
* Aleoholic.
Remarks—The fur of this bat is distributed as in Myotis yuman-
ensis yumanensis. On the back the distal portion of the fur is isabella
color and on the belly it is a pale tint of light buff. The skull differs
considerably from skulls of other races of Myotis yumanensis from
California. In addition to the greater size of the cranium, the brain-
case is more inflated, and at the same time the depression between the
brain-case and rostrum (in altipetens) is less marked than in the other
subspecies.
Transmitted April 14, 1916.
UNIVERSITY OF CALIFORNIA PUBLICATIONS
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Vol. 17, No. 3, pp. 11-14 September 18, 1916
Tas ES PES SE GN TE EEE Se EEN ST GE ESE Linmeeped 97
SPELERPES PLATYCEPHALUS, A NEW ALPINE
| SALAMANDER FROM THE YOSEMITE -
NATIONAL PARK, CALIFORNIA
BY
CHARLES LEWIS CAMP.
UNIVERSITY OF CALIFORNIA PRESS
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Vol. 17, No. 3, pp. 11-14 September 18, 1916
SPELERPES PLATYCEPHALUS, ANEW ALPINE
SALAMANDER FROM THE YOSEMITE
NATIONAL PARK, CALIFORNIA
BY
CHARLES LEWIS CAMP
(Contribution from the Museum of Vertebrate Zoology of the University of California)
One of the results of the Yosemite Natural History Survey recently
carried on under the auspices of the California Museum of Vertebrate
Zoology is the discovery of a new salamander from the high Sierra
Nevada. This alpine species is apparently most nearly related to
some of the forms of Spelerpes inhabiting the central Mexican plateau
and Mount Orizaba, and constitutes an interesting addition to the
Pacific fauna, Inasmuch as this genus has not hitherto been recorded
from west of New Mexico. The present species appears to be one of
a number of forms far removed geographically from the center of
abundance of the genus, and its locality of occurrence makes it seem
likely that the boreal salamanders of this group had a much wider
range during glacial times than at present. One is led to expect that
other species of Spelerpes may yet be found in western America, par-
ticularly in the higher regions of northern Mexico.
Spelerpes platycephalus, new species
Mount Lyell Salamander
Type—Female, adult, no. 5693, Mus. Vert. Zool.; head of Lyell
Canon, 10,800 feet altitude, Yosemite National Park, California;
July 18, 1915; collected by C. L. Camp; orig. no. 2215.
Diagnosis—Body elongate; tail shorter than head and body;
palatine teeth in two slightly arched series, separated from the para-
sphenoid patches and extending beyond the choanae; head broad and
12 University of California Publications in Zoology [ Vou. 17
depressed; tongue attached to central pedicel only, free in front;
digits half-webbed, 4-5; costal folds 12; color dark chocolate, mottled
on back, sides, limbs, feet, tail and chin with gray.
Material—Two adults from the type locality, as above: nos. 5693
female, and 5694 male, Mus. Vert. Zool.
Comparisons.—The free tongue, consolidated premaxillaries, ossi-
fied parietals, and digits 4-5 place the present species within the genus
Spelerpes. The half-webbed toes and character of the palatine teeth
relate it more closely to the Mexican alpine forms of that genus than
to those of the tropical and austroriparian regions of North America.
In number of costal folds and style of coloration (see Broechi, 1882,
pl. 19), it resembles S. leprosus Cope (1869, pp. 105-106), described
from the ‘‘ Alpine region, in Vera Cruz, Mexico.’’ But it is distinct
from leprosus because of its wider and longer head and shorter tail
(see table of measurements).
MEASUREMENTS IN MILLIMETERS OF SPELERPES
S. platycephalus
A
no. 5693 °
no. S. leprosus
in 5694 as
alive alcohol 3 (oh 9 6sex?2
MotallMleng Ghee cere eee 106.5 99.4 86.9 91 105 87
Tail from posterior angle of vent 35.5 33.6 30.2 45 52 37
lead! widthy fcc ncecccesceerccretse ese eee Haley) 10.3 10.2 8 9 8
Snout to gular folde=s 15.3 14.2 12.6 3 —_
Slows ose Ll yee eee ee 20.7 18.4 2 ats 15
STAG) Rea ON Seer ereeriereeersen Seca 54.2 47.8 ee ee 46
de) cil De oe eseeceeer eee a eee ee ete 16 15.5 15.5 12 14 15
OS Wes Ul hse 9 yes sec ee ecer cece: Heater 18 17.0 16.9 5) 17 18
* Boulenger, 1882, p. 68.
* Cope, 1869, p. 106.
Description of type-——Parasphenoid teeth in two elongate patches,
separated by a narrow interval from the palatine teeth which extend
in two slightly arched series beyond the choanae; head depressed,
wider than body at any point; no trace of canthus rostralis, subnarial
protuberances or parotoid glands; naso-labial grooves barely dis-
tinguishable ; nostrils minute; premaxillaries united, containing a fon-
tanel; maxillary and mandibular teeth small and numerous, jaws not
edentulous posteriorly ; body elongate, containing distance from snout
to gular fold four and one-half times; limbs fail to meet by two costal
interspaces, when appressed to sides; fore limb forward to middle of
eye; hind limb forward over five costal folds; fingers and toes slightly
1916] Camp: Spelerpes platycephalus 13
enlarged at ends, webbed about half way to tips, inner digit rudi-
mentary, smaller than outer on both front and hind foot; tail much
shorter than body, cylindrical; eyes small and far apart; extended
tongue 30-40 millimeters long; gular fold continuous with a line
running to eye; costal folds 12.
Color in life dark chocolate, marked evenly and thickly on back
and sides of head and body, on limbs, feet and tail, and below chin
with lichen-lhke gray markings, yellowish in tinge down middle of
back, where less distinct, and bluish on sides; underparts posterior to
gular fold uniform dark chocolate. Iris bright yellow. In alcohol
the yellowish tints have disappeared.
Fig. 1.—Top of head, 2 no. 5693, Mus. Vert. Zool.; note great breadth of
head, and distance between eyes.
Fig. 2—Side of head, 2 no. 5693, Mus. Vert. Zool.; note small nostril, rela-
tive size of eye, position of gular fold, and lines on side of neck.
Fig. 3.—Open mouth of ¢ no. 5694, Mus. Vert. Zool.; note tongue, unattached
in front, character of palatine and parasphenoid teeth, and long maxillary
teeth.
Fig. 4.—Left fore foot, J no. 5694, Mus. Vert. Zool.; note extent of webbing.
5
Fig. 5—Left hind foot, ¢ no. 5694, Mus. Vert. Zool.; note extent of webbing,
2)
and enlarged ends of toes. All x 2}.
Variations—The only other specimen, a male, differs somewhat
from the type. The teeth on the sides of the upper jaw (see fig. 3)
are abnormally long, protruding below the closed lips beneath the
eyes, and are few in number (ten on each side) ; the body is shorter,
the appressed limbs being separated by only one costal fold, and the
light markings are smaller and more scattered.
Remarks.—The two Mount Lyell salamanders were taken on the
rocky, snow-covered north slope of Mount Lyell, in the Yosemite
National Park, about a mile from the glacier and a little below timber
line, here marked by a few stunted white-bark pines on the tops of
14 University of California Publications in Zoology [ Vou. 17
the ridges. The exact spot was at the 10,800-foot contour, on a steep,
east-facing hillside above the Donohue Pass trail in a small patch of
heather. A stream close by issued directly from the snow banks and
disappeared beneath rock-slides below. The two specimens were found
to have been captured simultaneously in a spring-clip mouse-trap set
in front of a small hole running into the moist soil beneath some rocks.
Transmitted August 25, 1916.
LITERATURE CITED
BOULENGER, G. A.
1882. Catalogue of the Batrachia Gradientia s. Caudata and Batrachia
Apoda in the collection of the British Museum. (London, 1882),
second ed., pp. vili+127, 9 pls. :
Broccui, P.
1882. Mission scientifique au Mexique et dans 1’Amérique Centrale, ouvrage
publié par ordre du ministre de 1’instruction publique. Recherches
zoologiques. Troisiéme partie, 2° section, Etude des batraciens
de 1’Amérique Centrale. (Paris, 1882), 122 pp., 21 pls.
Copr, E. D.
1869. A review of the species of the Plethodontidae and Desmognathidae.
Proce. Acad. Nat. Sci. Phila., 1869, pp. 93-118.
os
0,1
gure in
aioe mek
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Vol. 17, No. 4, pp. 15-20, 1 figure in text October 3,\1916
A NEW SPERMOPHILE FROM THE SAN
JOAQUIN VALLEY, CALIFORNIA, WITH
NOTES ON AMMOSPERMOPHILUS
NELSONI NELSONI MERRIAM
BY
WALTER P. TAYLOR
(Contribution from the Museum of Vertebrate Zoology of the University of California)
Accumulation of specimens of antelope chipmunks (genus Ammo-
spermoplilus) from the San Joaquin Valley, California, is now
sufficient to permit a review of their systematic status, and this review
demonstrates the existence of a new subspecies, herewith described,
in the vicinity of Los Banos, Merced County. Occasion is taken also
to comment on the distribution of Ammospermophilus nelsoni nelsoni,
to make a contribution to its life-history, and to discuss its economic
relations.
The writer desires to thank Dr. Joseph Grinnell and Mr. Harry
S. Swarth, of the staff of the Museum of Vertebrate Zoology, for
helpful suggestions.
Ammospermophilus nelsoni amplus, new subspecies
Los Banos Antelope Chipmunk
Type—Male adult, in summer pelage; no. 16693, Mus. Vert. Zool. ;
20 miles south of Los Banos, Merced County, California; June 20,
1912; collected by R. H. Beck; orig. no. (J. Grinnell) 1957.
Diagnosis—Similar to Ammospermophilus nelsoni nelsoni but
larger, paler colored, and with white lateral lines less distinct.
Comparison—Summer skins of Ammospermophilus nelsoni amplus
are paler than those of A. n. nelson’. Their dorsal coloration approx-
imates light buff, almost whitish on nape of neck and sides of face,
16 University of California Publications in Zoology — | Vou. 17
while in nelsoni the hue is close to ight pinkish emnamon, approach-
ing whitish back of ears. A tendency is observable in amplus to have
the white of the lateral stripes a little obscured by an admixture of
light buffy hairs, particularly towards the rump. In nelsoni the
stripes are purer white. The general pallor of amplus also renders
the white stripes less conspicuous. The white eye-ring is also less
prominent in amplus than in nelsom. The pure coloration of the
upper surface of fore and hind lmbs tends toward pinkish buff in
amplus, more towards light pinkish cmnamon in nelsoni, although in
certain examples no difference in hue is appreciable. There seems
to be a tendency in amplus toward a flatter tail than in nelsoni.
The coloration of the tail seems also to be more clearly disposed in
bands than it is in nelsoni, im which the tail dorsally is more indis-
criminately mixed, blackish and whitish.
There are no conspicuous cranial differences between the two
forms. Certain crania of Ammospermophilus nelsoni amplus average
2.7 per cent longer than in A. n. nelsoni, mastoid width about 3.0
per cent greater, and nasals about 9.8 per cent longer.
Difference in size usually furnishes a comparatively conspicuous
character. Total length in Ammospermophilus nelsoni amplus aver-
ages 9.6 per cent above the average in A. n. nelsoni; length of tail
vertebrae, 5.4 per cent, length of hind foot, 6.0 per cent, and length
of ear, 40.8 per cent. Unfortunately the length of ear cannot be
measured, by the methods used, accurately enough to be as reliable
as are the other measurements.
With two exceptions the specimens of Ammospermophilus nelsoni
amplus are im summer pelage. Two examples (nos. 13816, 13817,
Mus. Vert. Zool.), taken March 25 and March 27 respectively, are
in winter coat. The pelage in winter is softer and thicker than in
summer, and is more grizzled or spotty; hues of coloration are paler,
and the white stripes on the sides less conspicuous. One of the
examples (no. 13816) is inclined more toward grayish or whitish;
the other (no. 13817) is more yellowish.
Distribution.—Of Ammospermophilus nelsoni amplus there are 34
specimens (including three skeletons-only) in the Museum of Verte-
brate Zoology, representing two localities, as follows: Sweeney’s
Ranch, 22 miles south of Los Banos, California, 2; mouth of Little
Panoche Creek, 18 or 20 miles south of Los Banos, 32.
Remarks.—Although Ammospermophilus nelsoni amplus is a well-
marked form, it is sufficiently close to A. n. nelsoni to be referred to
1916] Taylor: A New Spermophile from the San Joaquin Valley 17
the same species. According to present records there is a gap between
the ranges of the two from about the vicinity of Huron, Fresno
County, northward to a point within 18 or 20 miles of Los Banos,
Merced County; but it is not improbable that wide-spread collecting
in the region of this seeming gap would demonstrate continuous
distribution and geographic intergradation.
Measurements.—Of the type: Total length, 260 mm.; tail verte-
brae, 75; hind foot, 43; ear, 7; greatest length of skull, 43.1; mastoid
width, 21.6; interorbital constriction, 10.5; length of nasals, 13.1.
Males average slightly larger than females. External measurements
of 15 males: Total length, 249 mm. (max. 267, min. 234); tail
vertebrae, 73.1 (max. 78, min. 66) ; hind foot, 41.3 (max. 44, min. 40).
Corresponding measurements of 16 females: Total length, 238 (max.
256, min. 230) ; tail vertebrae, 72 (max. 78, min. 67); hind foot, 40
(maxe 43°) min: 31/9)
External measurements of 31 adults, combining both males and
females: total length, 243.6 mm. (max. 267. min. 230) ; tail vertebrae,
72.5 (max. 78, min. 66) ; hind foot, 40.6 (max. 44, min. 37).
Crania of four males: Greatest length of skull, 42.2 mm. (max.
43.1, min. 41.5); mastoid width, 21.4 (max. 21.6, min. 21.2); inter-
orbital constriction, 10.2. (max. 10.5, min. 9.9); length of nasals,
12.4 (max. 13.1, min. 12.2). Corresponding measurements of three
females: Greatest length of skull, 40.1 (only one specimen measured) ;
mastoid width, 21.1 (max. 21.7, min. 20.5, two specimens measured) ;
interorbital constriction, 9.7 (max. 10.2, min. 9.2); length of nasals,
12.0 (max. 12.5, min. 11.6, two specimens measured).
Cranial measurements of seven adults, combining both males and
females: Greatest length of skull, 41.8 (max. 43.1, min. 40.1);
mastoid width, 21.38 (max. 21.7, min. 20.5) ; interorbital constriction,
9.9 (max. 10.5, min. 9.2) ; length of nasals, 12.3 (max. 13.1, min. 11.6).
NOTES ON THE DISTRIBUTION AND LIFE HISTORY OF
AMMOSPERMOPHILUS NELSONI NELSONI
Distribution.—This species was first made known by Dr. C. Hart
Merriam in 1893 (pp. 129-131). That its distribution is general in
the southern San Joaquin Valley is indicated by the accompanying
map (see fig. 1). Actual records, as published by Merriam, pertain
to the following localities: Tipton (type locality), Tulare County ;
Alila (now known as Earlimart), Tulare County; Huron, Fresno
18 University of Califorma Publications in Zoology [Vou 1+
County; Adobe Station, Kern County (see Palmer, 1893, p. 362) ;
Lerdo, Kern County; Poso, Kern County (see Palmer, 1893, p. 378) ;
Lake Buena Vista, Kern County; Temblor (or Temploa) Mountains,
on boundary between Kern and San Luis Obispo counties. The
following additional stations of occurrence, ascertained by the Museum
of Vertebrate Zoology, may now be added: Eight miles northeast of
—
e A.n nelsoni
4 A.n. amplus
Fig. 1.—San Joaquin Valley, California, with outlines of ranges of Ammo-
spermophilus nelsoni amplus and Ammospermophilus nelsoni nelsoni. Dots
represent localities where specimens have been collected.
Bakersfield, Kern County; McKittrick, Kern County; and Carrizo
Plains, seven miles southeast of Simmler, San Luis Obispo County.
In 1912, Grinnell (MS) noted many antelope chipmunks in the
Cuyama Valley, extreme southeastern San Luis Obispo County.
General habits—Antelope chipmunks were found in abundance
by our Museum party eight miles northeast of Bakersfield, where
the animals preferred the open, exposed tracts covered chiefly with
foxtail grass. They were noted also in Atriplex country both in the
1916] Taylor: A New Spermophile from the San Joaquin Valley 19
vicinity of Bakersfield and in the neighborhood of McKittrick, where
sandy washes seemed to be fairly well occupied. Near Bakersfield the
antelope chipmunks were found in the hilly regions only, none at
all being noted on the plains below. They do not, however, so far
as available data indicate, range above the Lower Sonoran zone. Their
call-notes were not loud, being, in fact, indicated more by the associated
convulsive movement of the body than by the sound itself. During
the cool of the morning the animals were not in evidence, appearing
only after it became quite warm. Apparently six or eight individuals
constituted a colony of average size. The sides of little gullies were
often fairly honeycombed with burrows, such situations on banks being
preferred, perhaps, to any other location. The burrows investigated
proved to be somewhat complicated, two or three passageways running
into one at a depth of a foot or more below the surface. This passage-
way, in turn, joined one or two other aggregations of entrance ways.
Although several burrows were dug out to the end, no nests or food-
stores were discovered. The cheek pouch of a specimen from
McKittrick (no. 9032, Mus. Vert. Zool.), contained 745 seeds of
Erodium (either moschatum or cicutarium).
Breeding habits—Adult males (taken May 7 to 28) had testes
enlarged, signifying reproductive activity. Two juvenal specimens
were collected, a male (no. 13824, Mus. Vert. Zool.), May 7, 1911,
eight miles northeast of Bakersfield, and another (no. 13850, Mus.
Vert. Zool.), May 18, 1911, at McKittrick. No embryos were found,
so probably the young had been born during April or very early in
May, or perhaps even earlier.
Economic relations.—The Nelson chipmunk seems to prefer barren
situations whieh have no agricultural importance. The possibility
of its doing damage by burrowing through the walls of irrigating
ditches or by eating the seeds of cultivated plants is remote. The
experience of the Museum party, and of other observers with related
species (for example, see Mearns, 1907, pp. 300 and following),
supports the conclusion that the members of this group of spermophiles
(Ammospermophilus) are of little or no economic importance.
Possible effect of farming on the status of the species.—No antelope
chipmunks could be found in the vicinity of the type locality, Tipton,
Tulare County, the first noted being observed some 30 miles to the
south. Residents acquainted with the species testified that it formerly
occurred where now it is absent. Possibly farming activity crowds
out Ammospermophilus. Residents also asserted that the Beechey
20 University of California Publications in Zoology Vou. 17
ground squirrel is a comparatively recent immigrant into the lowlands
of the San Joaquin Valley, and that it is continually increasing in
numbers. Possibly the immigration of Citellus beecheyi tends to
assist in the crowding out of Ammospermophilus nelsoni and the
consequent restriction of the range of the latter. Much more evidence,
however, must be gathered before any degree of probability can be
assumed for the suggestions here made.
Transmitted April 21, 1916.
LITERATURE CITED
MEARNS, E. A.
1907. Mammals of the Mexican Boundary of the United States. U. 8S.
Nat. Mus. Bull. 56, Part 1, pp. xv +530, pls. 13, 126 figs. in text.
MerrIAM, C. H.
1893. Descriptions of eight new ground squirrels of the genera Spermo-
philus and Tamias from California, Texas, and Mexico. Proce. Biol.
Soe. Wash., 8, pp. 129-138.
PatmeEr, T. S.
1893. List of localities visited by the Death Valley expedition. U. 8.
Dept. Agric., Div. Ornith. and Mamm., N. Amer. Fauna, 7, pp.
361-384.
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UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 5, pp. 21-58, plates 1-4, 2 figures in text October 26, 1916
HABITS AND FOOD OF THE ROADRUNNER IN
CALIFORNIA
BY
HAROLD C. BRYANT
(Contribution from the Museum of Vertebrate Zoology of the University of California)
CONTENTS
Tintin OC Gi ony eee eee ene eee peat pec:
DMlaybe rial) es cecccscesce ets nacceeccccsccncctseecnce:
Life-history and habits ....................-
Food of the roadrunner in California
Vegetable food
Animal food
Kein Swamdquanbitvesy ots LOO dmb emis execs senescence ce er ran co see eeene nee a
Variation of food according to time of year
CS HO CEH i ELD] OE Oh ya eS a eee 43
THOOMONMANG, GHEYRUS) Cosceeteercescete cee reeset sepa ee eRe See eae a MER ne Re Ear ee ey 43
SHOU STEED ea tee eee eC een er 46
ERGOT UUM Om C1 GO Cpr enter cere te ee cre, Mes cece EE «ok noo doe 5 chsacdsasauacozuceestce st euseeeutecetseesees 48
INTRODUCTION
Investigations of the food habits of birds have of late done more
to afford birds proper protection than any discussion from a purely
esthetic point of view. It seems to the writer, however, that the
pendulum may have swung a little too far in the direction of the
dollars-and-cents view and that the esthetic is being left too far in
the background. Even to the economic ornithologist who is busy
determining the money value of birds there comes a vision of a day
when all true values will be taken into account and the monetary
faetor will be given only its due share of consideration. Nevertheless,
it is natural to emphasize the poimt of view which at the moment
makes the greatest appeal, and that at the present time appears to
be the economic.
22 University of California Publications in Zoology [Vou. 17
The esthetic worth of the roadrunner (Geococcyx californianus)
is great and should not be altogether overshadowed by other con-
siderations. Every early settler in California was quickly informed
of the paisano (‘‘the countryman’’), a bird which could run faster
than a horse, just as surely as he was informed of the ‘‘sacred toad’’
(horned toad), which wept tears of blood. The vaquero whose horse
was fast enough to overtake a roadrunner so that it could be lassoed
boasted of the fact for weeks. In fact, the roadrunner adds interest
and charm to every region where it is found. Recognition is, there-
fore, given to this and to other inherent values even though they be
not emphasized in this paper.
Because the roadrunner is conspicuous in size and action gunners
have used it as a mark. The added incentive to kill this bird because
of its alleged habit of destroying the eggs and young of the valley
quail, a favorite game bird, has been instrumental in practically ex-
terminating it in many localities in this state. Whether or not the
food habits of the roadrunner have justified this treatment has not
heretofore been adequately considered. The protection afforded the
bird in California has thus far rested wholly upon the few general
statements regarding its food habits and upon the recognition of
its esthetic value. The present investigation has been carried on
with the aim of providing a sounder basis either for its protection
as a valuable species or for its destruction as an injurious species.
MATERIAL
Kighty-four stomachs of roadrunners collected in southern Cali-
fornia, most of them in San Diego County, have been available for
examination. This material was obtained in 1911 and 1912, when
the California Fish and Game Commission was gathering information
as to the food habits of the western meadowlark. This paper is,
therefore, an additional report on the investigation then begun by
the Commission. The material embraced birds taken in every month
of the year with the exception of March. General complaint has been
made against the roadrunner that it is destructive of the eggs and
young of valley quail. Positive evidence on this point was sought
by collecting the largest number of birds during the nesting season
of quail. The deputies of the Fish and Game Commission who col-
lected the birds were instructed to take them in localities where quail
were nesting and where positive evidence as to their bird-eating habits
1916] Bryant: Habits and Food of the Roadrunner 23
could therefore be obtained. Deputy Webb Toms of San Diego, who
obtained the largest number of specimens, reported as follows: ‘‘I
commenced to collect roadrunners April 19, 1912, during which month
I obtained two. From May 1 to June 16, fifteen were procured. All
of those which I obtained during the nesting season were taken where
quail and other birds were nesting.’’ The entire number of birds
secured should suffice to furnish fairly complete evidence as to the
food habits of the roadrunner during the nesting season. The alleged
habit of destroying quail made it necessary to investigate the food
of the roadrunner in uncultivated districts as well as in those in which
the species comes in contact with civilization.
If the food habits of a bird found more abundantly in this state
were to be considered, a larger number of stomachs would be requisite
for examination, but the comparative rarity of the roadrunner did
not justify the killing of more individuals than seemed absolutely
necessary.
The material was handled and the same method (percentage-
volume) used as is described in a previous paper on the economic
status of the western meadowlark (Sturnella neglecta) in California
(H. C. Bryant, 1914, pp. 395-420). As in previous work, the attempt
has been made to furnish comparable data to all interested by using
a combination of the European and American methods. The tables
showing the number of birds taking the different items of food will
allow British economic ornithologists to compare readily the food
habits of the roadrunner with those of British birds. On the other
hand, the use of the percentage-volume method as developed by the
United States Biological Survey will furnish the data to which
economic ornithologists in America are accustomed.
My sincere thanks are due Miss Anna M. Lute of the Bureau of
Plant Industry, United States Department of Agriculture, for the
identification of seeds found in the stomachs. Professor Charles W.
Woodworth and Professor Edwin C. Van Dyke, of the Department
of Entomology in the University of California, have determined for
me, or verified the identification of, different species of insects, and
to them also I desire to make grateful acknowledgement. Mr. Donald
R. Dickey and Mr. William L. Finley kindly allowed the use of
excellent photographs, for plates 1 and 2. Mr. Tracy I. Storer,
Assistant Curator of Birds in the California Museum of Vertebrate
Zoology, has assisted in making photographs for illustration, and
Director Joseph Grinnell, of the same Museum, has put at my disposal
24 University of California Publications in Zoology [ Vou. 17
apparatus and information which has been of great help. He has
also critically read the manuseript and offered suggestions.
LIFE-HISTORY AND HABITS
Although most birds are found to possess at least some interesting
characteristics, there are very few which have so many outstanding
peculiarities as has the roadrunner (Geococcyx californianus). Four
marked characteristics distinguish this bird. First, although classified
with the cuckoos, it has no close relatives, other than the slightly
different Mexican roadrunner (Geococcyx affinis), living in North
America at the present time; second, it is probably the fastest endur-
ance runner among California birds; third, it has peculiar nesting
habits; and fourth, its diet is most extraordinary.
No better proof of the fact that the roadrunner has long excited
keen interest among people can be offered than that numerous stories
have grown up which attribute more than ordinary intelligence to
this bird. The commonest story, and one often implicitly believed,
is to the effect that the roadrunner has an ingenious method of killmg
a rattlesnake, a feat which it takes every opportunity to perform. This
myth has even crept into scientific writings (Heermann, 1859, p. 61;
Orcutt, 1886, p. 49; Van Dyke, 1897, pp. 36-88; Cooper, 1870, p.
369). The following version has been put on record by the orni-
thologist Walter E. Bryant (1891, p. 60):
It is said that when the roadrunner finds a rattlesnake coiled and asleep,
it corrals him or builds a fence around him of the cactus burrs with their innumer-
able sharp spines. After completing the corral it then commences to tease and
worry the snake by darting at and pecking it with its stout bill. The snake
in endeavoring to extricate himself from his thorny enclosure finds himself
pricked on every side by the sharp spines of the cactus, and tantalized by the
bird becomes infuriated, bites himself, and dies.
Furthermore, the many local names of the roadrunner show it
to be a well-known bird wherever found. In addition to the name
‘‘roadrunner’’, which is probably a translation of the Spanish name
correo del camino, it is variously called chaparral cock, ground cuckoo,
paisano (Spanish for ‘‘countryman’’), lizard bird, and snake bird.
Practically all of these names emphasize some peculiar characteristic
of this bird.
So unusual is the general appearance of the roadrunner that field
marks are readily employed as a means of identification. The legs
are long and powerful; the tail is as long as the body and rounded
1916 | Bryant: Habits and Food of the Roadrunner 25
at the end, and the outer tail feathers are tipped with white; the crest
when raised is conspicuous, as is also the bare skin about the eye,
which is red, blue, and whitish in color. The bill is over two inches long
and hooked at the tip. The plumage is conspicuously striped with
buffy brown and white and on the back and tail is glossed with green.
At close range it will be noted that two toes point forward and two
backward, a character which has caused this bird to be classified along
with the cuckoos. The character of the feet make the footprints along
a dusty road readily recognizable.
There is no bird in the arid southwest that is more characteristic
of the chaparral belt and desert. Most abundant in the San Diegan
and desert regions, the roadrunner becomes less numerous toward the
northern limit of its range in Shasta County. North of the Tehachapi
it is most common in the foothills of the Sierras. It has been re-
corded twice from the humid coast belt north of San Francisco Bay,
in Sonoma and Marin counties (Maillard, 1900, p. 63; Belding, 1890,
p. 56); and it approaches the coast in Alameda County (near Oak-
land) and in San Mateo and Santa Cruz counties. It has been found
rarely in the San Joaquin Valley. A few exceptions are noted: A
roadrunner was noted at Tranquillity, Fresno County, on June 22,
1915 (H. C. Bryant, MS). This individual sought refuge among tules,
an out-of-the-ordinary habitat for this bird. One was seen by Tyler
(1913, p. 82) at Lane’s Bridge, Fresno County, December 6, 1905,
and a few others near Fresno at other times. In the mountains of
southern California the roadrunner is occasionally seen above 5000
feet ; but at or above this altitude it is always on some hot slope where
Upper Sonoran vegetation is in evidence. Cooper (1870, p. 77)
reported the species as occurring on Catalina Island in the early 60’s,
but no additional records of its occurrence on the islands off the coast
of California are known. A study of the distribution of the road-
runner shows that the area inhabited follows very closely the limits
of the Upper and Lower Sonoran life-zones (Grinnell, 1907, pp.
51-53).
It is not obvious how the range of the species is affected by its
limited powers of flight, but the range of the individual is probably
comparatively extensive, due to its highly developed powers of pedal
locomotion. Where the individual range of most song-birds can be
computed as covering but a few acres, that of the roadrunner in all
probability needs to be computed in square miles.
The breeding season usually covers the months of March, April,
26 University of California Publications in Zoology [Vou. 17
and May. A nest placed in a cholla cactus and containing four eggs
was found by Kelsey (1903, p. 132) im Cholla Valley, San Diego
County, Mareh 25, 1903, and Sharp (1907, p. 87) found eggs advanced
in incubation as early as February at Escondido in the same county.
Another nest containing three nearly full-fledged young was discoy-
ered in a mesquite bush by Stephens (MS) at Salton, Imperial County,
as early as April 17, 1909; and a nest containing three fresh eggs was
found as late as July 16, 1904, by W. M. Pierce at Claremont, Los
Angeles County (Willett, 1912, p. 54). The foregoing instances in-
clude the earliest and latest nesting dates recorded for this state.
Although the roadrunner is largely terrestrial, it seeks safety
for eggs and young by placing its nest well above the ground. Some
thorny shrub or cactus is usually selected and the nest is placed from
three to ten feet above the ground (plates 1 and 2). Shepardson
(1915, p. 159) records having found one nest at an elevation of
twenty feet in a willow tree and another, in 1915, twelve feet above
the ground in a pepper tree growing in a well-populated section of
the city of Los Angeles. The nest itself is constructed of sticks loosely
laid together. A slight lining of finer material is to be found, almost
invariably composed of dry horse manure more or less broken up. A
cast-off (molted) snake skin, and feathers are also frequent adjuncts.
In approaching the nest the roadrunner usually ascends the cactus
or other shrub in which the nest is placed, by hopping from limb to
limb, seldom attempting to fly directly to the location.
The eggs number from three to nine and are deposited at intervals
of several days. Both male and female help during the incubation
period, which lasts about eighteen days (Bendire, 1895, p. 17). Incu-
bation is apparently begun when the first eggs are laid, for we have
considerable evidence that fresh eggs, small young, and fledglings are
found in the nest at the same time. Bendire (1878, p. 39) calls atten-
tion to this habit in the following words:
In 1872, while in southern Arizona, I found some twenty nests of Geococcyxr
californianus, the first nest on April 8, the last on September 10. During the
month of April, in which I found several nests, not one contained more than
three eggs, although I allowed incubation to begin before taking the eggs,
as I expected the birds to lay more. Nearly every nest I found after the
middle of May contained four or five eggs, and I account for the greater
number laid late in the season by the fact that insect food during the dry
season, which includes April and May, is comparatively scaree. The birds
being aware of this content themselves with rearing a small brood the first
time, and a large one at the second laying, when the young are hatched about
the beginning of the rainy season, which sets in in June. At this time all
1916 | Bryant: Habits and Food of the Roadrunner 27
kinds of insects and reptiles become exceedingly abundant, and the birds have
less trouble in providing for a family of five than earlier in the season for
one of three. Only occasionally have I found eggs in different stages of
incubation, and I do not believe that there was over a week’s difference in
the time of laying of the eggs in any nests I found.
No further evidence is at hand that more eggs are laid at a second
nesting later in the season. In fact, this seems rather improbable.
There is additional evidence, however, that eggs and young are often
found in the nest at the same time. Coues (1903, pp. 606-607) states
that perfectly fresh eggs and newly-hatched young may be found
together; and by the time the last young are breaking the shell the
others may be graded up to half the size of the adult. The Finleys
(1915, p. 162) found a nest in Arizona which contained one fresh
egg, one egg just ready to hatch, ‘‘two featherless, greasy, black
young,’’ and two young ones about ready to leave the nest. Moreom
(1887, p. 41) records the finding of a young roadrunner and two
eges In which incubation was advanced in a nest in San Gorgonio Pass,
Riverside County, California, on May 27, 1886.
”?
The young are described as ‘‘most hideous’’ in aspect. Their coal-
black skin and ungainly form combine to make them more or less
repulsive. When nestlings are disturbed they attempt to frighten
the intruder by making a clicking noise with their bills.
The habits of the roadrunner are of peculiar interest. The follow-
ing account to be found in the report of one of the early exploring
expeditions (Heermann, 1859, pp. 60-61) emphasizes its speed in
running:
We found this bird throughout California, frequenting at times the most
arid portions of the country. It often crossed our path, or ran before us for
a short distance on the road, dashing, when alarmed, immediately into the
chaparral, where, swift of foot, it easily evaded pursuit. It may, however, be
overtaken when followed on horseback over the vast open plains where no
friendly bush offers the weary bird a shelter. When closely chased, if on an
elevated point, it will sometimes fly, but always sailing downward. I once
saw one captured by a couple of dogs, their appetites whetted by recent suc-
cess in overtaking and bringing down a coyote or prairie wolf. Hotly pressed,
the bird would gain upon his enemies while sailing down the mountain slope,
but taking to his feet on the first ascent, this advantage was again soon
lost, and the fugitive, worn out, fell at length a victim to their relentless
determination.
H. W. Henshaw (1876, p. 257) states that the long tail of the
roadrunner, when running at full speed, is lowered till its end almost
touches the ground, and that the bird seems to fairly glide over the
28 University of California Publications in Zoology [ Vou. 17
earth, so easy are its movements. When hurrying, the tail is used
in turning quickly, being thrown from side to side with a jerk, accord-
ing to the direction to be taken. On gaining the cover of bushes, the
bird’s safety seems assured, and it usually pauses in the first cover
and stands with head erect, the tail vibrating nervously while it holds
itself in readiness to recommence its flight.
As a rule, this bird is shy and suspicious, but its inquisitiveness
sometimes so overcomes this character that it appears tame and con-
fiding. The species is not at all gregarious, no more than single indi-
viduals or pairs being ordinarily seen at one time.
Roadrunners kept in captivity by Sutton (1915, p. 58) refused
to bathe in water, but continually took dust baths. The operation is
described thus: ‘‘Shaking all over, and creeping along the ground
by flapping their wings inwardly, they make the dust fly into all
their feathers.’’
The small amount of water needed for sustenance probably
accounts for the facet that this species thrives on the hottest deserts
where the nearest water is several miles distant. In captivity the
roadrunner rarely drinks more than twice a week. When thirsty long
measured draughts are taken and the breast is dampened by the
proceeding (Sutton, 1915, p. 60).
Quick of action, the roadrunner is expert in catching its prey,
be it insect, lizard or mouse. When pursuing insects the bird will
often leap into the air eight or ten feet and catch them on the wing.
The commonest note of the roadrunner is a low ‘‘ br-r-r-r-r,’’ which
does not carry to any great distance. More often the only sound to
be heard from the bird is a snapping of the mandibles. During the
nesting season a sort of song much like the cooing of a dove is regu-
larly heard. This song is thus described by Holterhoff (1883, pp.
182-183) :
Standing near the summit of the hillock amidst his favorite cactus, and
with outstretched neck and head bent down, he would utter, as if by pro-
digious effort, the lugubrious notes I had wrongly thought the cooing of the
dove. At each iteration of the cry he seemed to make a renewed effort as
if to rid himself of the troublesome ‘‘whooo,’’ and when finished would stand
motionless, perhaps marveling at the sweetness of his own voice, or more
likely awaiting a response.
These peculiar notes of the roadrunner sound, as near as I can word it,
much like the prolonged syllable whooo—aspirating strongly the wh and giving
the vowel as a soft guttural. This note—not so prolonged as the ‘‘coo’’
of the dove—is repeated some five or six times in distinctly separate utter-
ances...
1916 | Bryant: Habits and Food of the Roadrunner 29
A roadrunner makes an amusing and interesting pet and in early
days in California this bird was often kept about the house and
garden and was used as a destroyer of insects and mice. Some inter-
esting accounts of these pets are to be found in literature.
FOOD OF THE ROADRUNNER IN CALIFORNIA
VEGETABLE Foop
About 10 per cent, 9.93 per cent to be exact, of the food of the
roadrunners examined was found to be made up of vegetable matter,
wild fruit and seeds being most in evidence. Unlke many birds which
turn their attention to vegetable food during the winter season, the
roadrunner appears to discriminate as to the kind of seeds taken.
The percentage volume of one particular kind of vegetable food was
found to be over ten times as great as of all others. This favorite
food is the fruit and seeds of the sour-berry (Rhus integrifolia), one
of the common sumaes of the southern part of the state. Twenty-
six birds, or 31.3 per cent of the stomachs examined, contained
the seeds or fruit of this shrub, and 8.4 per cent of the food taken
by all the birds was made up of this element. .The attention of the
roadrunner 1s apparently attracted to this vegetable food only during
the winter season, when insects, lizards, and other kinds of food are
least abundant.
No other fruit was found, and the only other seeds were atriplex
(Atriplex sp.), 0.01 per cent; cactus (Opuntia sp.), 0.35 per cent;
buckthorn (Rhamnus sp.), 0.01 per cent; and a few seeds of alfilaria
(Erodium sp.). Indeed, so small a percentage of seeds other than
the seeds of sumae and cactus were found that it may be that these
small seeds were taken into the stomach along with other food and
were not in reality chosen by the bird. The same can be said in regard
to the small rootlets and grass sometimes found in the stomachs, which
amounted to less than a tenth of 1 per cent. Attention has been called
(Bendire, 1895, p. 14) to the faet that the roadrunner eats the fruit
of cactus, but in the stomachs examined nothing but the seeds of the
opuntia was found.
Even though limited to desert and arid regions, a much wider
variety of fruit and seeds is available than is taken, as has been
shown by the results of stomach examination. The logical conclusion,
therefore, is that a preference has been established, by southern Cali-
fornia roadrunners at least, for the fruit and seeds of the sour-berry
(Rhus integrifolia).
30 University of Califorma Publications in Zoology [Vou. 17
ANIMAL Foop
The animal food of the roadrunner, which made up slightly over
90 per cent of the total food, is varied, perhaps as varied as that
of any common species of bird. Almost any animal, from the smaller
TABLE I
NuMBERS OF Foop ITEMS TAKEN BY FIFTEEN ROADRUNNERS DURING WINTER
Monrus, NOVEMBER, DECEMBER, JANUARY
n
Aa rneas fat . Bs 8 2d
= oh ae Dire, e 2 an os) % 3
te Ih ey Sse deh eh ee te
Bo on pt Gi ee See cele Roe temic ae eae
o Hal wu a & =I 355 o a = 5 ® AaI CI 4
ey ES 4s i te te fe) i im a GS ro El Si
January -....--..--. 17 10 =p
Ciao Ae | hoe eae Cy eee Ae 5 ? 1
2 1
1 8 Fane
baee ues BHOkey, 1
Oy 2% Wan 8 —
5 al
See 16 2 3
CC se er ake 4 ES 2
OS Wey se Dh py.8 28 oe
December .......... 1 > @ 4 fos
ce 9 pose == 4
oe > 3 1
[wees 1 il Ds tes 10 Lt
Chi 1B) 9 1 il
Rotalsaee S8e Miks 4509 4 So SN ew OWT A) G3 eal! 7
Average number of items per bird—13.6.
NuMBERS or Foop ITEMS TAKEN BY FIFTEEN ROADRUNNERS DURING SUMMER
MontTHS, JUNE, JULY, AUGUST
Be 3 8 E & «3 a «
o ° 2 i) 5 3 i S oe 5 2 s s
BOR io) 18) 0S Sy 3Bt Bee ah Sees
aye es op Sy tee SS) st dsl =) nee tS
2 3 =a 8 on
8 5 1 lie ae 1
1 4 iL
9 1 1
1 3 me
63 1 a8
Olea SF: il = 1
26 1 1
2 19 can
uy alta) 5 Z
6 1 ee
27 5 Secs
aes 5 1 1
2 4 2 250 1
Motals) 2. Tom ek ee eel 1 if 6 14 253 4 2
Average number of items per bird—36.2.
1916] Bryant: Habits and Food of the Roadrunner 31
rodents down to tiny insects, appears to be relished by this bird.
Although the stomachs examined showed no large percentage of verte-
brates, other published records show that reptiles sometimes form a
large part, if not the entire diet. Even these larger elements of food
are usually swallowed whole at one gulp. That the digestive appa-
ratus is powerful is evidenced by the fact that bone, hair, and feathers
pass through the digestive tract, and are not thrown back out through
the mouth in the form of pellets as is the ease with some hawks
and most owls.
Beetles—Beetles form next to the largest percentage of the animal
food of the roadrunner. Over 18 per cent (18.20 per cent) of all the
food of the roadrunners examined was made up of beetles of different
sorts. Since this bird is a ground-loving species, ground beetles of
various sorts are most often taken, more than sixty having been found
in a single stomach (see pl. 4, fig. 4). Such injurious beetles as the
click-beetles and wood-borers were also found in numbers in the
stomachs. Beetles are apparently taken in largest quantity during
the winter months, a total of eighty-eight having been found in the
food of fifteen birds during November, December, and January, as
against seventy-five in that of the same number of birds during June,
July, and August (see table 1). The following species of Coleoptera
were identified :
BEETLES (Coleoptera)
Amara californica Dej. Coniontis sp.
Amara insignis Dej. Coniontis elliptica (Casey)
Amara sp. Silpha ramosa Say
Eleodes sp. Scyphophorus yuccae Horn.
Lixus sp. (near pleuralis Rhynchitidae
Le Conte) Anisodactylus dilatatus De}.
Lixus sp. Crossidius intermedius Ulke
Eleodes acuticauda Le Conte Epicauda puneticollis Mann.
Miecroschatia inaequalis Le (?) Pterostichus sp.
Conte Asida sp.
Centrocleonus pilosus Le Conte Sphenophorus sp.
Grasshoppers and crickets——About one-third of the animal food
and almost one-fourth of the total food for the year was found to
be made up of grasshoppers (21.2 per cent). In addition, black
crickets (Gryllus sp.) made up 12.62 per cent and Jerusalem crickets
(Stenopelmatus) an additional 3 per cent, making Orthoptera a total
‘of 36.82 per cent of the food. Grasshoppers and crickets were taken
at all times of the year, but were most evident in the diet during
32 University of California Publications in Zoology [ Vou. 17
the summer and fall months (see table I). The largest number of
grasshoppers found in any one stomach was sixty-three. Many
stomachs contained between twenty-five and thirty. More individuals
had fed upon Orthoptera than upon any other group of insects (see
table IIL). Orthoptera, it can be seen therefore, make up the basic
element of the food of the roadrunner. The following species of
Orthoptera were identified :
GRASSHOPPERS AND CRICKETS (Orthoptera)
Mantidae Acridiidae
?Litaneutria pacifica Seud. Melanoplus differentialis Uhler
Gryllidae Melanoplus devastator Seud.
Gryllus pennsylvanicus Burm. Melanoplus sp.
Gryllus integer Seud. Schistocerca sp.
Locustidae ?Paropomala sp.
Stenopelmatus irregularis Brun.
Stenopelmatus sp.
Hollister (1908, p. 458) reports that he found the stomachs of
two roadrunners obtained near Needles, California, filled with ‘‘large
?
green grasshoppers.’’ A specimen taken by Stephens (MS) near
San Diego, November 5, 1908, was also found ‘‘crammed with grass-
hoppers.’’ In fact, so apparent has been the fact that the road-
runner feeds extensively on grasshoppers that practically every writer
has included these insects in statements regarding its food (see table
IL). So far as relative quantity per bird is concerned, the roadrunner
is probably to be numbered among the most efficient destroyers of
these familiar pests.
Butterflies and moths—The fact that butterflies and moths
usually escape attack by birds has again been demonstrated by this
investigation. Only a little over one-half of 1 per cent of the food
examined was made up of adult butterflies and moths. Two noctuid
moths were taken from one stomach, one from another, and a nymph-
alid butterfly from a third. Lepidopterous larvae, on the other hand,
formed 7 per cent of the food taken. One stomach contained as many
as thirty-six caterpillars. Three birds had taken woolly-bear cater-
pillars (Hustigmene acreae), another a white-lned sphinx caterpillar
(Celario lineata), and a third a pupa of Hemileuca sp. Hairy cater-
pillars of some species of Malacosoma were also found.
The roadrunner shows relationship with the cuckoos in its habit
of destroying hairy caterpillars. The stomach of a roadrunner taken
at Otay, San Diego County, April 19, 1912, contaimed seven large
3
«
Habits and Food of the Roadrunner
Bryant
1916]
Food OF THE
A. W. Anthony, 1896, p. 257; 1897, p. 217
ME Bailey 902) yi 0) eecesccseccssscrerecvecsesees
Baird, Brewer and Ridgway, 1874, p. 474
PHU OTCIT OLS O ony si ol duteenecane-saeecsnuece-cvesuseeceenene
Of, Wexennbingey, GSS Tob BI) aces creer eres rece bees
Wm. Brewster, 1883, p. 2
ee Gram COOP OL cL SMO sm psnre 09) meeeceeeene -ceeastenrssan
H. ‘Coues, 1903) pp: 606-607 -.2.222..e.
CO}, IDEN UES 105 PAVE) cerseccre enc coeeeoieeeren
\Vismilseee Ehas tre real! 0). sn 16 Oeemscsearesestaemsnrenesrteens
Grinnell, J., and Grinnell, H. W., 1907, p. 35
A. J. Grayson in W. HE. Bryant, 1891, p. 58
A. L. Heermann, 1859, pp. 60-61 ....
II. W. Henshaw, 1876, p. 257 .
N. Hollister, 1908, p. 458 ......... :
Knowlton & Ridgway, 1909, p. 449
D. E. Lantz, 1907, p. 51
F. Miller, 1879, p. 109
J. 8. Newberry, 1857, p. 51
C. H. Richardson, MS
R. W. Shufeldt, 1903, p. 177 ..
Cactus
Beetles
TABLE II
Grasshoppers
Crickets
ROADRUNNER ACCORDING
x &< Caterpillars
TO PUBLISHED REC
na
oo
S
a
Centipedes
Spiders
ks
wn
5
x)
A
as)
A
a
A
ORDS
Batrachians
Reptiles (not specified )
Mammals (not specified )
Rodents (not specified )
34 University of California Publications in Zoology [ Vou. 17
hairy caterpillars (Pseudohazis sp.), and at least four other birds
had taken from three to four hairy caterpillars apiece, but not of the
same species. One of the spiny type of caterpillar (Hwvanessa
antiopa) had been taken by a breeding female collected at El Toro,
Orange County, on May 18, 1912. One instance of the destruction of
caterpillars by the roadrunner, recorded by Anthony (1897, p. 217)
is of peculiar interest :
Not long since I called on a friend living in the suburbs of San Diego who
had a large number of unusually thrifty passion-vines climbing over his fence.
Upon inquiring the reason of their freedom from what I had considered an
inevitable pest, he informed me that a pair of roadrunners (Geococcyx cali-
fornianus) had for several months paid daily visits to his vines, climbing
through them in all directions until the last caterpillar (Agraulis vanillae)
had been captured.
Flies.—The only representatives of the order Diptera found were
two green-bottle flies (Lucilia caesar), a few robber flies and a syrphid
fly; these latter were undetermined as to genus or species. In addi-
tion, two stomachs contained quantities of fly larvae, those in one
stomach being probably larvae of some species of Musca. Hundreds
of these larvae were found in one stomach. The stomach of a road-
runner from the Imperial Valley contained the larva of a warble fly
(Cuterebra sp.).
Bugs.—F ive per cent of the total food was found to be made up
of hemipterous insects. Instead of a wide variety having been taken,
it was found that the birds examined had been largely choosing cicadas
(Cacama crepitans), insects which are close relatives of the seventeen-
year locust of the eastern United States. Cicadas formed 4.3 per cent
of the total food. Certain stomachs were completely filled with these
insects, as many as thirty-six having been taken from a single bird
(pl. 4, fig. 5). Eleven birds, or 13.2 per cent of all of those examined,
had fed upon this insect to some extent. The next most abundant
hemipteron was the cabbage-bug (Mergentia histrionica), ten having
been taken from a single stomach. The latter and other bugs
(Buschistus sp.; Anasa sp.) made up less than 1 per cent of the total
food.
Ants, bees, and wasps.—A little over 4 per cent of the total
food was made up of ants, bees, and wasps. Several bumblebees
(Bombus sp.), three carpenter bees (Ceritina sp.), and several
ichneumon flies were items taken by several birds. The stomach
of a roadrunner secured August 7, 1912, at Chula Vista, San Diego
1916] Bryant: Habits and Food of the Roadrunner 345)
County, contained over 250 red ants (Pogonomyrmex californicus),
along with a quantity of caterpillars, crickets, beetles, and grasshop-
pers. Another bird taken in October had eaten ten carpenter ants
(Camponotus sp.). One roadrunner had taken a tarantula hawk
(Pepsis sp.), and two had each taken a single honey bee (Apis
mellifera). The following species of hymenoptera were identified :
Ichneumonidae Apis mellifera Linn.
Bombus vornesenskii Rad. Chrysis sp.
Ceratina sp. Pogonomyrmex californicus Buck.
Pepsis sp. Camponotus sp.
Spiders and scorpions.—Spiders composed about 0.7 per cent of
the total food. A number of birds had taken the egg-cases of spiders
and one bird, taken August 1, 1911, at Sorrento, San Diego County,
had had the courage to devour a tarantula (Avicularia californica ?).
One of the outstanding features of the diet of the roadrunner in
California seems to be the presence of scorpions; these make up about
3.67 per cent of the total food. The commonest species found in the
stomachs was Anuroctonus phaeodactylus. Fourteen birds had taken
scorpions and the remains of as many as four were found in a single
stomach. Probably no other bird in California feeds so widely on
this particular arthropod.
Centipedes and millipedes—Both centipedes (Chilopoda) and
millipedes (Julus sp.) were found in the stomachs, but they formed
but a very small pereentage, much less than 1 per cent, of the total
food.
Crustaceans and snails—Sowbugs (Porcellio sp.) were found in
but one stomach. No further evidence that crustaceans were eaten
was forthcoming. McCall (in Baird, Brewer and Ridgeway, 1874, IT,
p. 474) states that in Texas snails are greedily eaten by the road-
runner: ‘‘These are usually taken either from the ground or a branch,
and carried to a particular spot, where the shell is broken and its
contents eaten. Piles of these shells are often found thus collected in
places frequented by them.’’ No snails were found in the stomachs
examined by the present writer.
Toads and frogs—Stomach examination gave no evidence that any
amphibian had been eaten. Toads were commonly eaten by tame
roadrunners kept by Sutton (1915, p. 59), but it is doubtful if these
vertebrates are taken by roadrunners under natural conditions.
Lizards and snakes—About 3.7 per cent of the total food was
found to be made up of reptiles. Except in one instance, lizards alone
36 University of California Publications in Zoology [ Vou. 17
had been taken. They were of the following species: Uta (Uta
stansburiana), blue-bellied lizard (Sceloporus biseriatus), and Blain-
ville horned toad (Phrynosoma blainvillet blainvillei). None of the
birds examined had eaten more than one lizard.
The following additional evidence that the roadrunner in Cali-
fornia feeds upon lizards is at hand: On July 9, 1894, near Pasadena
a roadrunner was secured, the stomach of which contained four full-
sized whip-tailed lizards (Grinnell, 1907, p. 85). C. H. Richardson
(MS) secured a roadrunner at Mecca, Riverside County, March 22,
1910, which contained, besides grasshoppers and beetles, a horned toad,
presumably of the desert species (Phrynosoma platyrhinos). The
stomach of a roadrunner taken at Needles, California, February 15,
1910, contained one weevil, remains of other beetles, and a half-grown
lizard (Cnemidophorus), the latter digested to the bare skeleton
(Grinnell, MS).
Finley and Finley (1915, p. 164) found that a roadrunner in
Arizona which they had under observation fed its young almost en-
tirely upon lizards. They write:
While we were crouching at the peephole of the blind the mother came,
carrying a big lizard, grasped firmly in her bill (plate 3). Up the bark
she scratched and thrust the lizard, head down, into the mouth of the
youngster. The tail hung out of its bill for a long time, but something
had hold of it down below, and finally it all disappeared. Soon she came with
another lizard, and presently another youngster was sitting propped stiffly
with a tail hanging out of its mouth. Again came a lizard—and again—and
again—there was no use counting. The larder was full of lizards and nothing
else!
Mr. Leo Wiley of Palo Verde, Imperial County, reports that he
has seen roadrunners carrying whip-tailed lizards and utas about,
although he has never actually seen the birds eat them. All evidence
points to the fact that the whip-tailed lizard (Cnemidophorus) is the
lizard most often eaten. Whether the abundance of this species is
responsible for this fact or whether a preference is indicated is not
apparent.
Very little attempt to break a lizard to pieces seems to be made
by the roadrunner. Instead, the reptile is usually swallowed whole
head first, after being hammered to death on some hard object nearby.
How the bird which had eaten the Blainville horned toad mentioned
above could have swallowed this reptile, which was fully an inch
wide, even when compressed, I am not able to say. But it appears
1916] Bryant: Habits and Food of the Roadrunner 37
that the throat of the bird must have power of expansion similar to
that exhibited by snakes.
The finding of a piece of the skin of a red-racer (Bascanian con-
strictor frenatum) in the stomach of a roadrunner taken at Palo
Verde, Imperial County, alone gave evidence that snakes form part.
of the diet. Although the present investigation furnishes but little
evidence that snakes are fed upon, yet the following records further
establish the fact that they are sometimes taken. A roadrunner ecap-
tured in New Mexico contained a garter snake a foot long (Bailey,
1902, p. 195); and Bendire (1895, p. 14) records the taking of a
garter snake twenty inches long from the stomach of a roadrunner
secured in Arizona. Mr. E. A. Goldman reported to me that the
stomach of a roadrunner which he secured in California contained
several young rattlesnakes. However, the idea that the roadrunner
is an active enemy of the rattlesnake is probably exaggerated. Rattle-
snakes may occasionally be eaten along with other snakes, but that they
are particularly sought out is extremely doubtful.
In this habit of feeding upon reptiles the roadrunner is almost
unique among birds, with perhaps the exception of certain hawks
and owls.
Birds.—Birds amounted to 1.7 per cent of the total food of the
roadrunners examined. The only evidence which showed that the
roadrunner feeds upon birds of any sort was that obtained through
the finding of the remains of a young Anthony towhee (Pipilo crissalis
senicula) and that of an Abert towhee (Pipilo aberti). No evidence
is at hand to determine whether these birds were taken alive from
the nest or secured after they had died. In the first instance the body
had been badly mutilated and the main parts left for identification
were the feet. The Abert towhee was taken from the stomach of
a roadrunner secured at Palo Verde, Imperial County. In this in-
stance the bird had been swallowed whole.
W. K. Fisher (1904, p. 80) states that an assistant of Professor
Leroy Abrams of the department of botany, Stanford University,
observed a roadrunner remove from a nest a young mockingbird and
devour it. Anthony (in Bendire, 1895, p. 14) says: ‘‘I know of
several instances of roadrunners making a meal of a nest of young
house finches (Carpodacus mexicanus frontalis) and other small
birds.’’ Beyond these instances we have no further information as
to the bird-eating habit among roadrunners, except the fact that birds
in captivity readily devour young sparrows or other birds furnished
38 University of Califorma Publications in Zoology [ Vou. 17
as food. A roadrunner kept at the State Game Farm at Hayward,
California, greedily devoured young sparrows and dead downy
pheasants.
Inquiries as to the destruction of young quail or eggs by the
roadrunner have elicited only the following information: Peter Lux,
of Encinitas, San Diego County, California, writes that he and his
brother saw a roadrunner catch and eat young quail at Olivenhain
about June 1, 1903. Mr. Lux also states that since that time (about
four years ago) he saw a young quail carried to a roadrunner’s nest
and fed to the young. Mr. George Wood, of Hollywood, California,
writes as follows:
In the month of June, 1906, I was making a study of the cause of the
death of so many young mockingbirds. I came on a nest containing two
young roadrunners, about two weeks old, and just at the moment of finding
the nest I also saw the male roadrunner with a young quail in its mouth.
It saw me and ran away. I got back under cover and watched until it
returned and I saw it take a young quail to the nest. J saw it return the
second time and repeat the above.
Mr. Leo Wiley of Palo Verde, Imperial County, recently told the
writer in a conversation that during the first part of August, 1915,
he saw a male roadrunner kill six Gambel quail which were about
four days old. The roadrunner gave each one a single blow with its
bill and, leaving the one just killed, attacked another.
Throughout the Imperial Valley it is rumored that the roadrunner
destroys hens’ eggs and young chicks. Mr. Walter E. Packard, in
charge of the Imperial Valley Agricultural Experiment Station,
writes: ‘‘T have heard a great many rumors to the effect that the
roadrunner sucks eggs, but I do not know of any instance where the
bird has been caught in the act or where it has been proven that
this bird was guilty of such an act. Nor can I refer you to anyone
who could give you information in this regard.’’ These accusations,
therefore, apparently rest on circumstantial evidence.
Such limited evidence as the above, unsupported as it is by the
results of stomach examination, would go to show that the habit of
destroying the eggs and young of birds is probably greatly exaggerated
and that the eggs and young of birds are only incidentally taken by
the roadrunner. Certainly the evidence at hand here in California
does not justify the wholesale destruction of the roadrunner on the
ground of its being an enemy of quail or other bird life.
Mammals.—Judging from stomach examination, 3.38 per cent of
1916 | Bryant: Habits and Food of the Roadrunner 39
the food of the roadrunner in California is made up of mammals.
Three different species were found in the stomachs: a harvest mouse
(Reithrodontomys megalotis longicaudus), a white-footed mouse
(Peromyscus sp.), and a newly-born San Diego cottontail rabbit (Syl-
vilagus auduboni sanctidieg?) .
Professor F. E. L. Beal procured a roadrunner in California which
had eaten a meadow mouse (Microtus californicus) (lantz, 1907, p.
51.) That even larger mammals are sometimes destroyed is evidenced
by the following observation made by A. W. Anthony (1896, pp.
257-258) :
This forenoon (May 7, 1896) I came suddenly upon a roadrunner (Geo-
coccyx californianus) that had just finished dispatching a wood rat (Neotoma).
The bird reluctantly withdrew as I came upon the scene, leaving the rat,
which I found to be quite dead. A post-morten disclosed a bad contusion on
the side directly over the heart, and another on the spine between the shoulders,
while the skull was crushed by a blow behind the ear, although the skin was
nowhere broken.
Ze
4
s S5
sco
Text Fig. A.—Diagram showing comparative amounts of the different kinds
of food taken by the roadrunner in California,
40 University of California Publications in Zoology [ Vou. 17
KINDS AND QUANTITIES OF FOOD ITEMS
The principal items of food as shown by this investigation and
the relative percentage volume of each are indicated in the accom-
panying figure (text fig. A). The outstanding feature is the
preponderance of insect food. The roadrunner is primarily in-
both vegetable and vertebrate food forming minor
elements. So far as bulk is concerned, the amount of grasshoppers
and crickets taken was double the amount of beetles and these elements
taken together formed more than half of the total food.
sectivorous,
TABLE IIL
NUMBERS AND PERCENTAGES OF ROADRUNNERS TAKING DIFFERENT KINDS OF FoopD
2
Zz iB 3
an o
2 : E pf 6E g 8
S 8 ¥ 2 5 Sn 5 S 2 ica} a
Ts = a S & = 2 = = a o a S
aes 3. E 2 8 Sa 2 & “ 2 =
Be of Bl « (Ae pale Re) See SS ee eee me
ae a = ° 3 n Bo mM 5S oS ‘S) a 4
26 2 il 5 1 4 3, 62 4 52 23 ait 4
$31.3 24 12 60 12 48 3.6 74.7 48 62:6 27.7 13.2 4:8
o
=
cs) zm
i = n 3
Sa Sis 3
a a oe a Sj w S 2 CS 5
Ba) eB ae ee et ee Sao 3
of 6§ 5 oe g 4 = 4E a = mI n g
eo P< 4 og a 8 o 12) = = 77) ~ Ls) 8
aS = & 25 2 Pe z S 8 = E et ia] S
Shs) A jay PeHH IS) q < Rn 1) = 1S) fan} f-2) =
15 4 2 5 11 22 8 14 2 2 2 4 2 4
18.2 48 24 6.0 13.2 265 7.2 168 24 24 24 48 24 6.0
A tabulation (table IIT) of the number of birds taking the differ-
ent elements of food is helpful in showing preference. It will be seen
that this table verifies the conclusions already made as a result of
the percentage-volume method. About three times as many birds
had taken beetles, grasshoppers and crickets as had taken any other
insect. Of the birds examined but four had taken reptiles, the same
number had taken mammals and but two had taken birds; the material
in one of these instances was doubtfully identified as being the intes-
tines of a bird. The fact that fourteen birds had taken scorpions
indicates that this invertebrate is a fairly common element of food
in the diet of the roadrunner. The table shows no other distinct
preference for a certain element of food.
1916 | Bryant: Habits and Food of the Roadrunner 41
PECULIARITIES OF THE DIET
The omnivorous food habits of the roadrunner have been amply
demonstrated by the above results of stomach examination, yet it is
desirable that some of the pecularities of the diet be emphasized.
As has already been stated, one of the outstanding features is the
apparent preference for one sort of vegetable food, the seeds and
fruit of a certaim sumac. Furthermore, the roadrunner apparently
turns its attention toward terrestrial vertebrates to a larger extent
than does any other California bird, with the exception of the raptorial
birds. As a destroyer of reptiles this bird probably takes first rank
among all the birds, and without doubt in many places in this state
the roadrunner is the worst natural enemy to which lizards and snakes
are exposed.
Many poisonous species of insects are eaten by the roadrunner.
The scorpion is perhaps the most conspicuous example. In many
instances the poison sae was found still attached to the tail of the
scorpion inside of the roadrunner’s stomach. No evidence was
obtained that ill effects are sustained by the bird from the poison.
Bumblebees come in the same category, as does also the single taran-
tula hawk taken. The latter is known to inflict a very painful sting.
If such insects as the latter were carefully killed by the bird before
being eaten, danger of being stung internally would be eliminated,
but the poison would still be present. A partial explanation of the
lack of ill effects may be that the poison is kept entirely within the
alimentary tract of the bird and has but little chance to enter directly
into the blood.
That the roadrunner has great ability in stalking rapid-flying
insects is illustrated by the fact that great quantities of cicadas are
consumed (see pl. 4, fig. 5). Entomological collectors state that cicadas
are difficult to capture. <A record of thirty-six cicadas in a single
stomach certainly shows that this bird is adept in securing certain
insects which other birds are not so successful in securing in quantity.
Probably the roadrunner relies upon its ability to stalk the imsects
rather than upon its speed on foot.
It is highly probable that the roadrunner destroys more hairy
caterpillars than does any other bird in the state of California.
Stomach examination has given abundant evidence that this bird preys
systematically upon these insects. The hairyness of a woolly-bear
caterpillar appears to be an effective protection to it in most instances,
but three of the roadrunners examined had eaten this insect. The
42 University of Califorma Publications in Zoology [ VoL. 17
California cuckoo, whose food habits are largely unknown, probably
displays this same habit of eating hairy caterpillars, as do other
cuckoos. But this species is so rare in California that its work in
comparison to that of the roadrunner must be slight.
The stomachs of two roadrunners each contained a praying mantis,
an insect seldom found in birds’ stomachs, perhaps because of its
comparative rarity. An ant lion (Myrmelion sp.) had been eaten
by one bird. A hairworm (?Gordius sp.) was found in one stomach,
but as this stomach also contained grasshoppers it is probable that
this parasitic worm originally may have been contained in one of
these insects.
VEGETABLE ANIMAL
JAN. 9
FEB. 2
MARCH o
APRIL 2
MAY 14
JUNE 8
JULY 12
AUG. 9
SEPT. 6
OcT. 6
NOV. 9
DEC. 6
Text-Fig. B.—Diagram showing the average percentage of vegetable food
(in white) and animal food (in black) consumed by the roadrunner in Cali-
fornia during the various months of the year. Numbers at the right indicate
the number of stomachs used in making the average.
VARIATION OF FOOD ACCORDING TO TIME OF YEAR
As is the case with most birds, the roadrunner is forced to vary
its food from one part of the year to another. In the winter when
insects become scarce vegetable food is resorted to. As can be seen
by a study of text-fig. B, but small amounts of vegetable food are
taken during the season when insects are abundant. A comparison
of the actual numbers of insects taken by fifteen roadrunners during
the summer months with those taken by the same number of birds
during the winter months (table I) shows that during the summer
season the roadrunner consumes on an average over twice as many
insects as it does during the winter months. No noteworthy seasonal
change in the kinds of insects taken is evidenced by the tabulation.
1916 | Bryant: Habits and Food of the Roadrunner 43
STOMACH CAPACITY
An average full stomach of a roadrunner contains about ten cubie
centimeters of food. The maximum quantity found in a stomach
was about forty cubic centimeters. This means that nearly two quarts
of food are needed weekly if the stomach were filled to an average
distension three times a day, and that a single bird probably consumes
in a year something like one hundred quarts. The fact that single
stomachs could contain over sixty ground beetles, sixty-three grass-
hoppers, or thirty-six large cicadas makes it evident that, so far as
the amount of food consumed is concerned, the roadrunner approaches
front rank among the common land birds. At first thought such a
capacity would seem to increase the ability of the bird to do good
or harm. This is in a measure true, but the much smaller number
of individuals of this species compared with certain gregarious species
of birds makes it possible that certain birds with much smaller stomach
capacity, because of the greater number of individuals, may consume
even more. The stomach capacity of the individual roadrunner is
great, but that of the total roadrunner population is relatively small.
Variety in the diet, as well as typical elements of the food, is well
illustrated by the following tabulation of the stomach contents of a
single roadrunner taken at Lemon Grove, San Diego County, January
2k, UGH} e
VEGETABLE, 3%
2 seeds (Rhus integrifolia) 2 grass-blades
1 seed with stem (Hrodium sp.)
ANIMAL, 97%
6 beetles (Amara insignis) 1 bug (Huschistus conspersus)
1 beetle (Hleodes acuticauda) 8 bugs (? Anasa sp.)
5 beetles (Microschatia inaequalis) 1 scorpion (Anuroctonus phaiodac-
5 beetles (Centrocleonus pilosus) _tylus)
8 wireworms (Elateridae) 1 spider egg-case
12 grasshoppers (Melanoplus sp.) 1 lizard (Uta stansburiana)
1 Jerusalem ericket (Stenopelmatus
sp.)
ECONOMIC STATUS
The investigation of the stomach contents of eighty-three road-
runners taken in southern California revealed but little evidence that
the roadrunner is injurious to man’s interests. The following, all of
minor importance, form the most important evidence against the bird:
Parts of an Anthony towhee found in one stomach; a very young
44 University of California Publications in Zoology [ Vou. 17
San Diego cottontail rabbit found in one stomach; three lizards, con-
sidered beneficial as insect destroyers, taken by three different birds;
and two honey bees found in different stomachs. Certain of the
carabid beetles found might by some be considered sufficiently bene-
ficial to warrant their preservation, but little is yet known of the
food habits of even the commonest ground-beetles destroyed by the
roadrunner, namely, the different species of Amara. Until more is
definitely known it cannot be said that the destruction of these beetles
by birds is distinctly detrimental.
The above adverse evidence is minimized in most instances by the
demonstrated fact that these elements of food are out-of-the-ordinary,
rather than staple items, for they form very small percentages of the
food for the year. This is not true, however, in regard to lizards,
for they form a staple article of diet. Should it be proved that lizards
are strictly beneficial as insect destroyers, this habit of feeding on
lizards might, according to present standards, serve to incriminate the
roadrunner. However, until conclusive evidence shows that the road-
runner actually reduces in numbers the scaly denizens of the desert
and in so doing is an adverse factor to inseet control, we are not
justified in assuming that the lizard-eating propensities of this bird
place a serious blot on its character.
The stories regarding the bird-eating habits of the roadrunner
appear to have some foundation in fact, but much of the evidence
submitted has been of the circumstantial kind, and with little doubt
some statements have been grossly exaggerated. The frequent asser-
tion that roadrunners commonly destroy the eggs and young of the
valley quail has not been substantiated by this investigation. No
evidence whatever has been obtained that they destroy the eggs of
quail or those of any other bird. That they do sometimes destroy young
quail and other young birds appears to be true, but statements to
the effect that such destruction is carried on systematically or that
young birds form a staple article of diet have not been verified.
Furthermore, our evidence fails to show that quail suffer destruction
to a greater extent than do other birds. In any ease, the bird-eating
habit appears to be individual rather than general.
Evidence pointing toward the usefulness of the roadrunner as a
valuable destroyer of insects and possibly of rodents, wherever it
comes in contact with civilization, was obtained as a result of the
investigation. Chief among the items of food the destruction of which
must be considered a benefit are: Grasshoppers, crickets, Jerusalem
1916 | Bryant: Habits and Food of the Roadrunner 45
erickets, wireworms, and hairy caterpillars. The regular consumption
of such large numbers of grasshoppers by roadrunners must in some
measure affect the abundance of the insect. The increase or decrease
in the numbers taken by the roadrunner according to whether the
insect is scarce or plentiful makes it probable that this bird, wherever
it is found, acts as one of the checks on grasshopper abundance.
* and black
crickets, although of less economic importance, are classified as in-
Jerusalem crickets, commonly known as ‘‘potato bugs,’
jurious insects. Wireworms are important insect pests in cultivated
fields, and their destruction by natural enemies is to be encouraged
rather than discouraged. Artificial means of destroying wireworms
are limited and the work of birds is therefore more potent. As there
are practically no vertebrate enemies of hairy caterpillars, and since
to this group of caterpillars belong many destructive larvae, their
systematic destruction by the roadrunner goes far towards establish-
ing it among those birds to be classed as beneficial. Add the possible
benefit conferred through the destruction of small rodent pests, and
the evidence in favor of protecting the bird exceeds the evidence
against it.
Economic ornithologists have many times called attention to the
fact that so far as apparent results are concerned the work of birds
in destroying insects is not so effective as an artificial method such
as the use of an insecticide. True it is, also, that birds less often
attack those injurious insects for the control of which large amounts
of money are spent than those which must be classified as of minor
importance. For instance, here in California, according to statistics
compiled by the Agricultural Experiment Station, about nineteen
times as much money is spent each year on the control of black and
other scales, phylloxera and codling moth, as is spent on all other
insects combined. Hence the commonest insects eaten by birds—
grasshoppers, cutworms and caterpillars, and wireworms—are of much
less economic importance and their destruction by birds appears to
be of less significance. Nevertheless, many of the workings of nature
are still beyond our knowledge and we are constantly finding out that
artificial methods are usually inferior to natural ones. May it not
be that the very reason why such insects as grasshoppers, cutworms,
and wireworms are of minor importance is that they are better con-
trolled by natural agencies? Certainly most of the insects of minor
economic importance are potentially as destructive as those for the
control of which great sums of money are spent. Indeed, the con-
46 University of Califorma Publications in Zoology [ Vou. 17
stant attempts to find and introduce some natural enemy of the more
injurious insects is proof that where the natural control is lacking
the most energy must be expended towards filling the need.
It must be granted that the roadrunner, like all others under
natural conditions, helps to maintain the balance of nature. And
even if its work in the destruction of economically important insects
is apparently of a less degree of importance, it should be remembered
that the bird may be performing a service which we cannot measure,
and that the systematic destruction of any of the natural enemies of
insects may be instrumental in increasing the economic importance
of that insect.
It is only when the roadrunner comes in contact with settled
communities that it can be said to be injurious, unless it be proved that
the number of quail accessible to the hunter is materially reduced.
Since most of the roadrunners are confined to desert regions far from
the haunts of men, the total harm done is much smaller than would
be the case with species more abundant in settled country. Then,
too, both the possible damage and the potential good are greatly
minimized by the fact that there is no concentration of individuals at
any one time or place. The number of roadrunners in any one
locality does not vary greatly and is always relatively small. Probably
the maximum numbers per square mile do not exceed ten or twelve
even in favorable localities.
SUMMARY
The roadrunner (Geococcyx californianus) is unique in many ways.
Of particular interest are its breeding habits and especially food
habits. It is doubtful if an investigation of the food habits of any
other American bird could have yielded such unusual and interesting
results.
The investigation here reported upon included the analysis of
eighty-four stomachs of roadrunners taken in southern California in
1911 and 1912 and also a survey of all available literature relative to
the food of the roadrunner in California. The stomachs became
available as a result of the inquiry into the food habits of non-game
birds instituted by the California Fish and Game Commission in 1911.
The eighty-four stomachs represented birds taken every month of
the year with the exception of March. The analysis of the stomach
contents showed that practically 90 per cent (90.07 per cent) of the
1916 | Bryant: Habits and Food of the Roadrunner 47
total food was made up of animal matter and that slightly less than
10 per cent was of vegetable material. Nearly all of the vegetable
matter was of one kind, the fruit and seeds of the sour-berry (Rhus
integrifolia). Insects and certain vertebrates composed the animal
food. Chief among the insects found were beetles (18.2 per cent),
grasshoppers and crickets (36.82 per cent), cutworms and cater-
pillars (7 per cent), cicadas and other hemipterous insects (5 per
cent), ants, bees, and wasps (4.24 per cent), and scorpions (3.67
per cent). Lizards of three species (3.73 per cent), two birds (1.56
per cent), one tiny cottontail rabbit (1.0 per cent), and two wild mice
of two different species (2.38 per cent) composed the vertebrate food.
The results of stomach examinations substantiated rather than
altered published statements regarding the food of the roadrunner.
From published sources, however, came added information as to the
number of snakes and lizards consumed by this bird and practically
all of the information regarding its bird-eating habits. One hzard,
the whip-tailed lizard (Cnemidophorus), appears to be taken more
often than any other species.
The individual capacity for food in this bird is great, for an
average full stomach contains about ten cubic centimeters of food.
The collective capacity, however, is small, due to the paucity of indi-
viduals. The amount of damage possible (and this must be said also
of the potential good) is greatly minimized because there is no con-
centration of individuals in any one place. A wide variety of food
items from small insects to reptiles and mammals is consumed.
Little evidence was obtained that the roadrunner is detrimental
to man’s interests. The destruction of a few beneficial insects and
birds, and of certain lizards usually considered beneficial, can alone
be taken as evidence against it. Even if the consumption of a certain
proportion of all of the above as food be a fixed habit, the end result
is minimized when the facts are taken into consideration that the
roadrunner is a bird of the desert regions rather than of the cultivated
fields and that the species exists in but small numbers.
A preponderance of evidence favors the bird. The destruction of
such unquestioned pests as grasshoppers, cutworms, caterpillars, and
wireworms, and of such rodents as mice is to be desired even if the
amount of destruction be relatively small. The taking of this sort
of food on wild land is evidence that this bird when feeding in
cultivated fields is likely to be distinctly beneficial.
Evidently the roadrunner never turns its attention to any sort
48 University of California Publications in Zoology [ Vou. 17
of cultivated crops as do those birds with granivorous and frugivorous
habits. On the contrary, a study of its food habits shows that the
roadrunner feeds upon the insect and rodent pests which attack cul-
tivated crops, and furthermore that it destroys hairy caterpillars, a
pest not commonly attacked by other birds.
As shown by this investigation, the food habits of the roadrunner
in California do not prove the bird to be harmful to any great extent.
Instead, it has been clearly shown that the benefits conferred through
the destruction of injurious insects and small rodents, though not
great, outweigh in value the harm done through the destruction of
lizards and birds. In fact, the destruction of a roadrunner can be
justified only under exceptional circumstances, and certainly the
wholesale destruction of the species because of its supposed habit of
destroying the eggs and young of quail is unwarranted. It is to the
interest of every citizen of the state to protect carefully this most
curious and interesting bird.
Transmitted March 22, 1916.
LITERATURE CITED
ANTHONY, A. W.
1896. The roadrunner as a rat-killer. Auk, 13, 257-258.
1897. The roadrunner as a destroyer of caterpillars. Auk, 14, 217.
BaiLey, F. M.
1902. Handbook of birds of the Western United States (Houghton-Mifflin
Co., N. Y.), xe +514, 36 pls., 601 figs. in text.
Bairp, S. F., BREwrr, T. M., and Ripeway, R.
1874. A history of North American birds. Land birds (Little, Brown &
Co., Boston), 2, 590+vi, pls. 27-56, many figs. in text.
BELDING, L.
1879. A partial list of the birds of central California. Proce. U. S. Nat.
Mus, 1, 388-449.
1890. Land birds of the Pacific district. Occ. Papers Calif. Acad. Sci., 2,
1-274.
BENDIRE, C.
1878. Breeding habits of Geococcyx californianus. Bull. Nutt. Orn. Club,
3, 39.
1895. lLife-histories of North American birds. Smithson. Inst. Contrib.
Knowl., 32, ix + 518, 7 pls.
BREWSTER, W.
1883. On a collection of birds lately made by Mr. F. Stephens in Arizona.
Bull. Nutt. Orn. Club, 8, 21-36.
BRYANT, H. C.
1914. A determination of the economic status of the western meadow-
lark (Sturnella neglecta) in California. Univ. Calif. Publ. Zool.,
11, pp. 377-510, pls. 21-24, 5 figs. in text.
1916] Bryant: Habits and Food of the Roadrunner 49
Bryant, W. E.
1891. Andrew Jackson Grayson. Zoe, 2, 34-68.
Cooper, J. G.
1870. Ornithology of California. Geol. Surv. Calif., vol. 1, Ornithology.
Published by authority of the Legislature (Univ. Press, Cam-
bridge), xi+592, many figs. in text.
Cougs, E.
1903. Key to North American birds (ed. 5; Dana Estes Co., Boston), 2,
vi+ 537-1152, figs. 354-747.
Davig, O.
1889. Nests and eggs of North American birds (ed. 4; Hann and Adair,
Columbus, O.), 10+455+ xii, 13 pls.
EVERMANN, B. W.
1881. California bird notes. Orn. & Ool., 6, 7-8.
Finuey, W. L. and FIN ey, I.
1915. With the Arizona roadrunners. Bird Lore, 17, 159-165, 7 figs. in text.
FisHer, W. K.
1904. Roadrunners eat young mockingbirds. Condor, 6, 80.
GRINNELL, J.
1898. Birds of the Pacifie slope of Los Angeles County. Publ. Pasadena
Acad. Sci., 2, 1-52.
1907. The California distribution of the roadrunner (Geococcyx califor-
nianus). Condor, 9, 51-53, 1 map.
GRINNELL, J., and GRINNELL, H. W.
1907. Reptiles of Los Angeles County, California. Throop Inst. Bull. 35,
1-64, 23 figs. in text.
HEERMANN, A. L.
1859. Report upon birds collected on the survey. Pae. R.R. Rep., 10,
Part IV, No. 2, 29-80, pls. 2-10.
HENSHAW, H. W.
1876. Report on the ornithology of the portions of California visited
during the field-season of 1875, by H. W. Henshaw. Ann. Rep.
Geog. Surv., W. 100th Merid., Append. H8 in Append. JJ, pp. 224—
278.
HOouuister, N.
1908. Birds of the region about Needles, California. Auk, 25, 455-462,
1 pl.
HOLTERHOFF, G., JR.
1881. A collector’s notes on the breeding of a few western birds. Amer.
Nat., 15, 208-219.
1883. Geococcyx as a vocalist. Bull. Nutt. Orn. Club, 8, 182-183.
KELSEY, F. W.
1903. The home of the California roadrunner. Condor, 5, 132-133, 2 figs.
in text.
MAILLIARD, J.
1900. Land birds of Marin County, Cal. Condor, 2, 62-68.
MILER, F.
1879. Strange story of a California bird. Bull. Nutt. Orn. Club, 4, 109-
110.
50 University of California Publications in Zoology [ Vo. 17
KNowttTon, F. A., and Rieway, R.
1909. Birds of the world (Holt & Co., N. Y.), xiv +873, 16 colored pls., 235
figs. in text.
Lantz, D. E.
1907. An economic study of field mice. U. 8. Dept. Agric., Biol. Surv.
Bull., 31, 1-64, 8 pls., 3 figs. in text.
Morcom, G. F.
1887. Notes on the birds of southern California and southwestern Arizona.
Bull. Ridg. Orn. Club, 2, pp. 36-57.
NEWBERRY, J. S.
1857. Report upon the zoology of the route. Report upon the birds.
Pac. R.R. Rep., 6, Part IV, No. 2, chap. 2, 73-110, 2 pls.
Orcurt, C. R.
1886. Roadrunners corralling rattlesnakes. West. Amer. Sci., 2, 49.
SHarp, C. S.
1907. The breeding birds of Escondido. Condor, 9, 84-91.
SHEPARDSON, D. I.
1915. Some western birds—roadrunner. Oologist, 32, 158-160.
SHUFELDT, R. W.
1903. The chaparral cock. Western Field, 2, 174-179, 2 figs. in text.
Surron, G. M.
1915. Suggestive methods of bird-study; pet roadrunners. Bird Lore, 17,
57-61, 1 fig. in text.
TYLER, J. G.
1913. Some birds of the Fresno district, California. Pac. Coast Avifauna,
9, 1-114.
Van Dykes, T. S.
1897. The roadrunner and the snake. Avifauna (Santa Barbara, Calif.),
1, no. 3, 36-38.
WILLETT, G.
1912. Birds of the Pacific Slope of southern California. Pac. Coast Avi-
fauna, 7, 1-122.
ug a a ”
i
me:
x
.
un :
i , :
‘1
i
; 1 an
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PLATE 1
Fig. 1—Nest of roadrunner situated in tangle of chilicothe and opuntia.
Photograph by Donald R. Dickey, taken at Lakeside, San Diego County, Cali-
fornia.
[52]
PLATE 2
Fig. 2—Roadrunner on nest in cactus. Photograph by Donald R. Dickey,
taken near San Diego, California.
[54]
UNIV, CALIF. PUBL, ZOOL. VOL, !7 [BRYANT] PLATE 2
PLATE 3
Fig. 3—Roadrunner carrying a whip-tailed lizard (Cnemidophorus). Photo-
graph taken by W. L. Finley near Tucson, Arizona. After Finley and Finley,
1915, by permission.
[56]
UNIV, CALIF. PUBL. ZOOL. VOL. 17 [BRYANT] PLATE 3
: ne
tee
Ly ae oy
i i ais
PLATE 4
Fig. 4——Stomach contents of a roadrunner taken at Nestor, San Diego
County, January 19, 1913. The stomach contained twenty-four beetles (Micro-
schatia inaequalis), one large beetle (EHleodes sp.), one beetle (Amara sp.),
one wireworm, and thirteen Jerusalem crickets (Stenopelmatus sp.).
Fig. 5.—Parts of thirty-six cicadas (Cacama crepitans) found in the stomach
of a roadrunner taken at Otay, San Diego County, May 30, 1912. The stomach
also contained one carabid beetle, four short-winged grasshoppers, one robber
fly, one stink bug, one carpenter bee, and one small spider.
[58]
UNIV. CALIF. PUBL. ZOOL. VOL. 17 [BRYANT] PLATE 4
o ibe FBB8
in text
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4, An Account of the Mammals and Birds of the Lower Colorado Valley, 7
with Especial Reference to the Distributienal Problems Presented,
by Joseph Grinnell, Pp. 51-294, plates 3-13, 9 text figs. March, 1914. 2.40.
5. Aplodontia chryseola, a New Mountain Beaver from the Trinity Region -
of Northern California, by Louise Kellogg. Pp. 295-296.
6. A Previously Undescribed Aplodontia from the Middie worth Coast of
California, by Walter P. Taylor. Pp.-297-300.
Nos..6 and 6 in one cover.) April, 19U4 uci. cecetennateswereernees 08
7. A Second Species of the Mammalian Genus Microdipodops from Cali- .. —
fornia, by Joseph Grinnell. Pp, 301-304, April, 1914 OB
8. Distribution of River Otters in California, with Description of a New. ~
Subspecies, by Joseph Grinnell. Pp. 305-310, plate.14. October, 1914 . .05
9, Four New Pocket Gophers from Cao: by Joseph Grinnell. EP.
$11-316.- ‘November, 1914 2 ew ee “(05
10. Three New Races of Vespertilionid Bats. from California, by Hilda
$2 a8
ZAS
GA ROARS een bi
Bia ad eta F pate at Ta
Wood Grinnell, Pp. 317-321. December, 1914 22.2 OB
11. Eutamias sonomae, a-New Chipmunk from the Tuner Northern Coast
Belt of ee by Joseph Grinnell. gous 321-325, 1 text figure.
TODAY; TOG aes ane Se et er ete ee I ti
UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 6, pp. 59-62, 4 figures in text November 17, 1916
DESCRIPTION OF BUFO CANORUS, A NEW
TOAD FROM THE YOSEMITE
NATIONAL PARK
BY
CHARLES LEWIS CAMP
(Contribution from the Museum of Vertebrate Zoology of the University of California)
Included in the material gathered by the Yosemite Natural History
Survey, and now being worked up at the Museum of Vertebrate Zool-
ogy under the direction of Dr. Joseph Grinnell, are a number of toads
of a species which up to the present time has apparently escaped the
attention of naturalists. Comparison with a large series of toads of
the boreas group at hand shows that the new species possesses char-
acters not included within the limits of that variable assemblage.
Bufo canorus, new species
Yosemite Park Toad
Type.—FKemale, adult ; no. 5744, Mus. Vert. Zool.; Poreupine Flat,
8100 feet altitude, Yosemite National Park, Mariposa County, Cali-
fornia; July 1, 1915; collected by C. L. Camp; orig. no. 2129.
Diagnosis—A medium sized toad with no head crests; parotoids
large, flat, circular, separated by a space usually less than their own
diameter and never greater; vertebral streak wanting in most males
and represented in other specimens by a thread-like, white line. Color
distinctive, and differing greatly in the two sexes; females with irreg-
ular, dark blotches, each usually bordered narrowly with white; males
speckled with black on a dull greenish background and without dis-
tinet dark patches on back.
60 University of California Publications in Zoology — | Vou. 17
Material—F orty-two alcoholic specimens, all but one from within
the Yosemite National Park at the following localities: Near Peregoy
Meadow, 7000 feet altitude (no. 57429) ; near Mono Meadow, 7300 feet
(no. 57474) ; East Fork Indian Cafion, near B. M. 7937 feet (no.
57282) ; vicinity of Porcupine Flat, 8100-8200 feet (nos. 57299, 57309,
57369, 57439, 57449, 57592) ; vicinity of Ragged Peak, 9700—10,200
feet (nos. 5721-5727, all ¢¢, 57319, 57329, 57349, 57359, 57373,
57384) ; head of Lyell Cafion, 9700-11,000 feet (nos. 57382, 5739,
57409, 574101, 57452, 57489, 5749-5753, all Jog, 5754-5758, all 99,
5760¢) ; Vogelsang Lake, 10,350 feet (mos. 603592, 60449). One speci-
men from Tioga Lake, Mono County, 10,000 feet (no. 57462).
Bufo canorus; 9 type; all X 1
Fig. 1. Top of head; note great width of parotoids and shape of snout.
Fig. 2. Side of head; note extent of parotoids and steep profile of snout.
Fig. 3. Front foot, from below; note small inner metacarpal tubercle.
Fig. 4. Hind foot, from below; note reduced webbing and large outer meta-
tarsal tubercle.
Comparisons.—In profile, lack of head crests, small tympanum, and
short legs this toad resembles Bufo boreas and its subspecies, but may
be distinguished at once from these forms by its smaller size, enormous
width of parotoids, slight interval between parotoids, very smooth skin,
absence of a broad vertebral stripe, and markedly different color pat-
tern in both sexes. In extent of webbing of hind foot the present
species most nearly resembles B. boreas halophilus, its near neighbor
in the southern Sierra Nevada and the San Joaquin Valley. Speci-
mens of B. boreas boreas from Mono County, directly to the east of
the range of canorus, have the large hind foot characteristic of the
more northern subspecies.
1916 | Camp: Description of Bufo canorus 61
MEASUREMENTS IN MILLIMETERS OF TyYPR SPECIMEN
Total length TIN Cy iO Ot Berens sseeee eee eeee es 39.0
Head length 18.9 Spread of hind foot —...... 22.8
Head width _.. . 26.0 Length of parotoid ........... 12.2
Hind leg - 82.0 Width of parotoid —........ 10.7
DUG ah too 2 en csssmeeueesevccsevsaecavesesss 2OLO Interorbital space .............. 7.2
Color (in alcohol). —Adult female, no. 5744, the type: All dorsal
and lateral surfaces, including sides of head, rostrum and parotoids,
thickly marbled with irregular but clearly defined, dark patches, each
surrounded by a brilliant white line; many of the dark patches end
abruptly at the mid-line, and do not cross it except in the anal region ;
ground-eolor almost white, becoming brownish on the back and paro-
toids; large tubercles in centers of dark patches tipped with brown;
limbs mottled like rest of body; underparts clear white except for a
few obscure dark specks; skin between large tubercles very smooth ;
no trace of speeckling or of small dots between the dark blotches. This
coloration is typical of all of the females, even the smallest.
Adult male, no. 5747, typical in coloration of all the males in the
series: Dorsad and laterad the body and limbs are covered evenly with
minute dots of black on a uniform olive-green ground-color ; each dark
speck surrounded by a narrow white ring; underparts grayish white
with scattered dots of a larger size than those on the back. The skin
is exceptionally smooth, and the larger tubercles and even the paro-
toids are scarcely discernible.
Remarks.—This toad is extraordinary in its pronounced sexual
dimorphism. The striking black and white pattern of the female is
entirely lost in most of the males, but a few of the latter show, by
traces of blotching about the head and legs, that their speckled pattern
is the result of a breaking up into smaller dots of the larger blotches
of the female. In coloration the female of Bufo canorus bears some
resemblance to the brighter patterns of Bufo boreas; the male phase
has some counterpart in the latter species in the case of an occasional
male of boreas, which assumes a dull green color and smooth skin
during the breeding season. In the series of canorus the glandular
development in the males is much less than in the females; in most
of the males even the parotoids are not well developed, though their
characteristic circular outline ean still be traced.
The web of the hind foot is small; the spread of the hind foot is
about 30 per cent of the total body-length, as in Bufo boreas halophilus.
The sole is free from the asperities developed by toads of more arid
62 University of California Publications in Zoology — [Vou. 17
regions, and both inner and outer metatarsal tubercles are distinct
and nearly equal in size. The inner metacarpal tubercle is less than
half the size of the outer.
Distribution and habitat—The limits of the range of Bufo canorus
north and south of the Yosemite Park are unknown. The species
seems to be absent from the high altitudes of Placer County and from
the Sierras south of the Kings River, where its place is taken by
Bufo boreas halophilus, which there goes to a high elevation. It is
the only toad found in the higher parts of the Yosemite National Park,
where it inhabits wet meadows and lake shores from about 7000 feet
altitude on Bridal Veil Creek among lodge-pole pines to 11,000 feet on
the headwaters of the Lyell Fork of the Tuolumne River, near the
upper limit of the white bark pines. It does not seem to live at the
lower altitude of the Yosemite Valley proper, where the larger species.
Bufo boreas halophilus, is found.
The specific name selected, canorus, refers to the long-sustained,
melodious trill uttered by this toad. This diurnal singing accompanies
the breeding activities, which take place as soon as the snow melts
from the Sierran meadows, June 1 to July 15. Many of the females
captured at this time contained mature eggs.
Transmitted October 5, 1916.
UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
i ZOOLOGY
Vol. 17, No. 7, pp. 63-74 December 28, 1916
7 5
THE SUBSPECIES OF SCELOPORUS Beet
OCCIDENTALIS
WITH DESCRIPTION OF A NEW FORM FROM THE
SIERRA NEVADA AND SYSTEMATIC NOTES
ON OTHER CALIFORNIA LIZARDS
BY
CHARLES LEWIS CAMP
UNIVERSITY OF CALIFORNIA PRESS
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with Especial Reference to the Distributional Problems Presented,
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5. Aplodontia chryseola, a New Mountain Beaver from the Trinity Region
of Northern California, by Louise Kellogg. Pp. 295-296.
6. A Previously Undescribed Aplodontia from the Middie North Coast of
California, by Walter P. Taylor. Pp. 297-300.
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Subspecies, by Joseph Grinnell. Pp. 305-310, plate 14. October, 1914. .05
9, Four New Pocket Gophers from California, by. Joseph Grinnell. Pp,
S11-S16. November, 1914 oo nk ot a eee .. 05
10. Three New Races of Vespertilionid Bats from California, by Hilda
Wood Grinnell, Pp. 317-321. December, 1914 2: Bt) 5)
11, Eutamias sonomae, a New Chipmunk from the Inner Northern ‘Coast
Belt of California, by Joseph Grinnell. Pp. 321-325, 1 text figure.
ATURE YO To a a Eh ee Oe.
12, Batrachoseps major and Bufo cognatus californicus, New Amphibia
from Southern California, by Charles Lewis Camp. - Pp. 327-334.
April “1915305 Se ee ee
Be Saat Nee 55 |
no VE a
- ee ae Seen
UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 7, pp. 63-74 December 28, 1916
THE SUBSPECIES OF SCELOPORUS
OCCIDENTALIS
WITH DESCRIPTION OF A NEW FORM FROM THE SIERRA
NEVADA AND SYSTEMATIC NOTES ON OTHER
CALIFORNIA LIZARDS
BY
CHARLES LEWIS CAMP
(Contribution from the Museum of Vertebrate Zoology of the University of California)
CONTENTS
PAGE
INGLES OUR SCElOPOLUSOCCICON DAIS es. .cse cere ohe cele econ sence se eet gc eee cee oe Be 63
Sceloporus occidentalis taylori, new subspecies ...............2...1-221:.0-22seseeese-eeeeeeeeeee- 66
HID TESMOTIES COLOWO GUM OM AG) OSUS tee seta see cnces aac cae hen cee vaca vee san ecove te oegae sense sneer an cedeeeeoee Gi
Sceloporus graciosus vandenburgianus Cope ............-......-----.--21.-----+------ sek 67
NOLES MOMMU Lele SUNS DUNN Aa cee tess tects = co cts-5 J. cecsc0islesessacensoteesstesies kaevsse. suse eet 68
Notes on Callisaurus and Crotaphytus - 70
IGLOS PMO MMR ON EMIT ODN OLUS WEL OTIS) eee ee. ccc eeancan thc necont accanenencosnccaveesnnien se dees net oeenae eaeoewonae 71
WarteGlonmis Or sHMIMeCes (SKUCOULATMUS: soo ccc. e cee we acne cease Seraceenene seer mer see YE)
NOTRE EEE ASS MOAI CG LS oe Fe 3 eRe ee eae a ee 74
Noves ON SCELOPORUS OCCIDENTALIS
In a systematic treatment of the lizards of the genus Sceloporus
occurring in the Pacifie district, Van Denburgh (1897, pp. 73-74,
77-84) calls the two closely related forms, occidentalis and bi-seriatus.
distinct species, and separates them by the characters of size and of
the degree of confluence of the throat patches in males. An exam-
ination of the series of about 900 specimens representing these two
64 University of California Publications in Zoology | Vou. 17
forms, contained in the Museum of Vertebrate Zoology, has revealed
the existence of a definite intergradation between them through an
extensive area in California.
In males from points within the range of typical occidentalis, in
the Sacramento Valley and in Humboldt, Marin, Napa and Alameda
counties, about 72 per cent have the throat patches joined as in all
male bi-seriatus. The other 28 per cent of the males is what we should
have to rely on for the separation of these races if we held to this
character alone. Our series of occidentalis from eastern Mendocino
County averages about as large in size as a series of small bi-seriatus
from Los Angeles County. It therefore becomes desirable to seek out
some additional character as a criterion for the separation of these
two forms, if they are, indeed, to be considered worthy of systematic
notice at all.
Nearly every male occidentalis I have examined can be distin-
euished from bi-seriatus by the greater amount of light color on the
underparts, particularly on the lower surface of the hind limbs and
on the chest and between the belly patches. A single specimen of
occidentalis from Monterey and ten or so of a large series from the
coast of northern California have the parts mentioned dark greenish
or slaty in color. A male from Bakersfield, and another from Little
Rock Creek, Los Angeles County, each has an unspotted ght stripe
down the center of the belly and cannot be distinguished from average
occidentalis, although obtained well within the range of bi-seriatus.
But these are exceptions in a very large series which shows the gen-
eral duskier color of the subspecies bi-seriatus.
The most interesting thing about this character of the color of the
underparts is the distribution of the intergrades, which occupy a zone
running from the coast near San Luis Obispo northeast to western
Merced County, thence across the San Joaquin Valley to Coulterville,
Mariposa County, then northward over the Sierras to eastern Modoe
County. The range of occidentalis northwest of this zone contrasts
in its greater humidity with the range of bi-seriatus to the southeast
of it.
In the intergrades the white chest, mid-ventral line and hind limbs
are flecked by numerous dark scales, either widely separated (as in
occidentalis), or jomed into a general dusky suffusion as in bi-seriatus.
The body-length of individuals increases noticeably as one goes south
through the Sacramento and San Joaquin valleys from Yolo and
Solano counties into Fresno and Kern counties. This increase in size
1
1916 ] Camp: Subspecies of Sceloporus occidentalis 6!
o
and change in coloration of the underparts is most abrupt in the nar-
row district lying between the Merced River and Raymond, Madera
County. Adult males from San Emigdio and Walker Basin, in Kern
County, are from 75 to 90 millimeters in length of body (averaging 85
millimeters). The largest males of occidentalis from various localities
in the Sacramento Valley and in Modoe and Humboldt counties range
from 62 to 80 millimeters and average about 71. Those from such
intermediate points as eastern Merced County, and Raymond, Madera
County, are from 71 to 80 millimeters long and average about 75.
A key that will apply invariably for all individuals of both sexes
of the following three subspecies cannot be formulated. Females
without blue throat patches can be referred to S. 0. occidentalis, those
with a single throat patch to S. 0. bi-servatus (see Van Denburgh, 1897,
p. 73), but beyond this the identification of females is best not at-
tempted. Males (told at once by the enlarged post-anal plates) can be
distinguished by the following key:
A. Belly-patches separate from throat patches and divided by either a lighter
or darker band.
b. Chest, mid-ventral line and lower surface of hind limb very light
in color or speckled with darker scales; blue throat patches in
young males sometimes divided by light or dark seales; body
length of adults 62-80 millimeters.
Sceloporus occidentalis occidentalis Baird and Girard,
b’. Chest, mid-ventral line and lower surface of hind limb grey or
black in color; blue throat patch never divided; body length
of adults 75-90 millimeters.
Sceloporus occidentalis bi-seriatus Hallowell.
A’. Belly-patches confluent with throat patch and not divided by a lighter or
darker band; ventral color deep blue throughout; little or no dusky
color on chest.
Sceloporus occidentalis taylori Camp.
It is desired here to describe a very distinct new subspecies of
Sceloporus occidentalis occurring about the headwaters of the Merced
River in the Yosemite National Park and perhaps farther south in the
Sierra Nevada. Specimens of this brilliantly colored lizard were col-
lected by various members of the field party from the Museum of
Vertebrate Zoology, which participated in the Natural History Survey
of the Yosemite region during 1914-1915.
66 University of California Publications in Zoology — (Vou. 17
Sceloporus occidentalis taylori, new subspecies
Tenaya Blue-bellied Lizard
Type.—Male, adult; no. 5947, Mus. Vert. Zool.; half way between
Merced Lake and Sunrise Trail (Eeho Creek basin), altitude 7500 feet,
Yosemite National Park, California; August 25, 1915; collected by
Walter P. Taylor; orig. no. 7361.
Diagnosis.—Size, equalling the largest Sceloporus occidentalis bi-
seriatus ; underparts, in the male, blue throughout; belly-patches not
separated by a lighter or darker mid-ventral line; throat evenly
colored, heht blue to snout and lips, and lighter in tint than general
ventral color; blue of belly not separated from throat patch by a
hehter or darker area across gular region (young specimens excepted).
Female more richly colored below than in bi-seriatus; lighter than
male; belly-patches separated by a faintly lighter area; chest lehter
than belly; one extensive throat patch as in male; blue not always
extending to beneath hind limb.
Color (in alcohol) —Belly alizarine blue (of Ridgway, 1912), in
darkest males, to clear cadet blue in the lightest females; throat diva
blue to light cadet blue; chest only slightly dusky in the darkest speci-
mens; males with hind limbs beneath and anterior border of anus,
greenish blue, nearly as dark as belly. Back very dark as in darkest
bi-seriatus ; sides and some seales on back greenish; lighter and darker
ereseentie markings on back obscure, mest so in males. Females with
four series of small light spots down back.
Material—Fourteen males and ten females from the higher eleva-
tions of the Yosemite National Park at the following localities: Wash-
burn Lake, 7640-7700 feet ; near Merced Lake, 7500 feet; Echo Creek
basin, Merced River, 7300-7500 feet; lower McClure Fork, Merced
River, 7800 feet; Lake Tenaya, 8100 feet; and Glen Aulin, Tuolumne
River, 7300 feet.
Remarks.
A number of individuals at hand in a large series of
S. 0. bi-seriatus from the southern Sierras in Kern and Tulare counties
and farther north are, of all our specimens, the closest in size and
ventral coloration to taylori; they are, however, of greenish and more
dusky shades of blue beneath than the new form, and their status
must be held questionable pending the acquisition of material from
the headwaters of the Kings and San Joaquin rivers. A male speci-
men, one of two, from the Yosemite Valley, 4000 feet altitude, seems
1916] Camp: Subspecies of Sceloporus occidentalis 67
to be intermediate in size and color between this form and a series of
S. 0. occidentalis at hand from western Mariposa County. In the
Yosemite Park this subspecies lives on rocky, sunlit slopes in the
heavily glaciated region in the upper Merced basin, about Lake
Tenaya, and in the head of the Tuolumne Canon, at altitudes between
7300 and 8200 feet.
NOTES ON SCELOPORUS GRACIOSUS
The representatives of Sceloporus graciosus isolated upon several
of the mountain ranges of southern California south of Mount Pinos,
Ventura County, appear to constitute a locally differentiated race. It
is here proposed to reinstate the name vandenburgianus (Cope, 1896,
p. 834) for this subspecies, the type locality of which is the ‘‘ Summit
of the Coast Range, San Diego County, Calif.’’—probably in the
yellow pine belt on the Laguna Mountains, San Diego County.
Sceloporus graciosus vandenburgianus Cope
Van Denburgh Lizard
Diagnosis.—hLike S. g. graciosus, but adult males with blue belly-
patches united across mid-line, or else separated by a very narrow
interval only; belly-patches confluent with throat patch; undersurface
of hind limb and tail dark blue; the lighter, broad, mid-ventral line
and chest-patch of graciosus, indefinite or absent; blue of underparts
sometimes nearly black, usually darker than in graciosus. Females
more dusky in color beneath than females of graciosus.
Material—Ninety-two specimens of Sceloporus graciosus vanden-
burgianus from the following localities in southern California: San
Diego County: a single adult male from the Cuyamaca Mountains, 6000
feet altitude; Riverside County: Santa Rosa Peak, 7500 feet, Santa
Rosa Mountains, 7; San Jacinto Mountains, 21, from Thomas Moun-
tain, Strawberry Valley (6000 feet), Tahquitz Peak (8000 feet), near
Round Valley (8500 feet), vicinity of Fuller’s Mill (5850-7000 feet),
and vicinity of Schain’s Ranch (5200-5500 feet); San Bernardino
Mountains, San Bernardino County: 32, from Fish Creek (6500-6700
feet), South Fork of Santa Ana River (6200 feet), Santa Ana River
(5500 feet), and Clark Hill (6000 feet) ; San Gabriel Mountains, Los
Angeles County: 31, from vieinity of Mount Wilson (4150-5800 feet),
Mount Lowe (4000 feet), and vicinity of Pine Flats (5500-5800 feet).
68 University of Califorma Publications in Zoology [ Vou. 17
Of Sceloporus graciosus graciosus there are in the Museum of Verte-
brate Zoology 282 specimens, from Mount Pinos, Ventura County, the
southern Sierra Nevada, central and northern California, Humboldt
County, Nevada, and Wallula, Washington.
Remarlis—The series of sixty-three vandenburgianus from the San
Gabriel and San Bernardino mountains is nearer graciosus in colora-
tion than specimens from farther southeast. Several large males (nos.
776, 777, 781, 783, 790) from the San Bernardino Mountains eannot
be told from typical vandenburgianus, and some others are lighter ven-
trally than any males from south of San Gorgonio Pass. The series
from the San Gabriel Mountains includes the lightest-colored males
of the subspecies, and they are in this respeet very good intermediates,
despite their isolated station, between the small, hght-colored graciosus
of Mount Pinos and the large, dark blue specimens of vandenburgianus
from the headwaters of the Santa Ana River, and farther south.
Cases of this sort of distribution bear on the question whether
intermediates between two adjacent forms may be hybrids. In the
present instance the low passes of Soledad, Cajon and San Gorgonio
separate the range of the intermediates into a chain of mountain-top
‘islands’? and seem to preclude the possibility of hybridization at the
present time.
The darker ventral color and larger size of the southern race of
this species is interestingly paralleled in Sceloporus occidentalis, Ger-
rhonotus scincicauda, Cnemidophorus tigris, and Eumeces skillonianus,
in which species the southern and desert forms are bluer or blacker
beneath and duskier above.
Novres on UTA STANSBURIANA
Richardson (1915, pp. 412-418) separates Uta stansburiana into
the following three subspecies: U. s. stansburiana, characterized by
its greater number (average 103.4) of dorsal rows of weakly carinated
seales and its few femoral pores (average 13 +); U. s. elegans, with
from 78 to 103 (average 86) dorsal rows of more heavily keeled scales,
an average of 14-+ femoral pores and a greater tail length; and U. s.
hesperis, with from 87 to 117 dorsal rows of heavily carinated scales,
and an average of 14+ femoral pores. He extends the range of the
typical subspecies, sfansburiana, into California on the basis of two
specimens from Round Valley and Lone Pine, Inyo County. Counts
of dorsal scale rows of some additional specimens of this species in
1916 | Camp: Subspecies of Sceloporus occidentalis 69
the Museum of Vertebrate Zoology from Inyo County seem to indi-
cate that western Inyo County is the locality in California where
intergradation between slansburiana and elegans takes place. Table I
shows how the number of scale rows increases as one goes north
through Owens Valley.
I. TABLE SHOWING NUMBERS OF DORSAL TRANSVERSE SCALE ROWS IN SPECIMENS
OF UTA S. STANSBURIANA AND U. S. ELEGANS
Mus. Dorsal seale
No. Locality rows Remarks
3105 Mohave, Kern Co. 83 elegans
3644 Keeler, Inyo Co. 87 elegans
3104 Carroll Creek, Inyo Co. 97 intergrade
Average of two specimens, Independence,
Inyo Co. 99 intergrade
Average of five specimens, Kearsarge
Pass, at 6000 feet, Inyo Co. 99 intergrade
6072 Laws, Inyo Co. 93 intergrade
Average of three specimens, Benton,
Mono Co. 102 stansburiana
Average of fifty-five specimens,* Utah,
Nevada, Idaho, and Oregon 103 stansburiana
* According to Richardson (1915, p. 418).
Counts of seale rows in twenty specimens of stansburiana picked
at random from series collected at a number of points in the San
Joaquin Valley and in San Luis Obispo County indicate that the
subspecies occurring in this region is hesperis, rather than elegans,
as Richardson (1915, p. 414) has indieated. The intergradation be-
tween hesperis and elegans seems to occur at such places as northern
Los Angeles County, Walker, Cajon and San Gorgonio passes, and
eastern San Diego County. This is shown in table IT.
II. Taste SHowtnc NuMBER OF DORSAL TRANSVERSE SCALE Rows IN SPECIMENS
OF UTA S. ELEGANS AND U. S. HESPERIS
Mus. Dorsal seale
No. Locality rows Remarks
3099 Kelso Valley, Kern Co. 86 elegans
3102 Fay Creek, near Weldon, Kern Co. 95 intergrade
3075 West side Walker Pass, Kern Co. 93 intergrade
3084 West side Walker Pass, Kern Co. 92 intergrade
Average of twenty specimens from San
Joaquin Valley and San Luis Obispo Co. 100 hesperis
885 Gorman, Los Angeles Co. 93 intergrade
4175 Pallett, Los Angeles Co. 94 intergrade
603 Cajon Wash, San Bernardino Co, 97 intergrade
0
605 Cajon Wash, San Bernardino Co. lt intergrade
70 University of California Publications in Zoology — | Vou. 17
Mus. Dorsal seale
No. Locality rows Remarks
5396 Victorville, San Bernardino Co. 77 elegans
5397 Victorville, San Bernardino Co. 79 elegans
3580 Banning, Riverside Co. 91 intergrade
90 Snow Creek, Riverside Co. or intergrade
252 Palm Canon, Riverside Co. 97 intergrade
491 Carrizo Creek, San Diego Co. 88 elegans
1043 Warner Pass, San Diego Co. 86 elegans
1585 Julian, San Diego Co. 90 intergrade
986 Mountain Spring, San Diego Co. 91 intergrade
3587 Imperial Valley, Imperial Co. 77 elegans
NovTEs ON CALLISAURUS AND CROTAPHYTUS
In his description of Callisaurus ventralis myurus, Richardson
(1915, p. 410) gives the range of this new form, doubtfully, as ex-
tending as far south as Owens Valley, in California. The darkest of
ten specimens in the Museum of Vertebrate Zoology from Keeler, Inyo
County, are not different in color from individuals collected at Barstow
on the Mojave desert in San Bernardino County. The average number
of femoral pores in this series is 14.3 and the average ratio of tail
to body length .737, thus agreeing with Richardson’s determination
of .727 for C. v. ventralis from the Colorado desert, and differing from
the average of .807 given for myurus from Pyramid Lake, Nevada.
Since Callisaurus does not seem to range north through Owens Valley
into Mono County, specimens from the former region are not in what
could be considered an area of intergradation between ventralis and
MYUrUs.
It should be noted that the provisional assignment of the Death
Valley Callisaurus to myurus on the basis of Merriam’s notes is a
mistake, since the Callisawrus of which Merriam speaks pertained to
Desert Valley, east of the Pahroe Mountains, Nevada (Stejneger, 1893,
p. LT).
It is interesting to add that Richardson’s discovery of a decrease
in tail length and number of femoral pores in the more northern forms
of widely ranging species of lizards can be confirmed upon a study
of Crotaphytus collaris baileyi and Crotaphytus wislizenii. Ten ex-
amples of the former species in the Museum of Vertebrate Zoology
from Humboldt County, Nevada, have an average ratio of body length
to tail length of .549 (minimum .521) and an average of 16 + (maxi-
mum 19) femoral pores. Eight specimens of baileyi from southern
California have an average ratio of body length to tail length of 476
1916 | Camp: Subspecies of Sceloporus occidentalis fall
(maximum .491) and femoral pores averaging 20+ on each thigh
(minimum 18). Three specimens from the Inyo Mountains and the
Kern River, five miles above Kernyille, are .450, .517, and .530, in
body-tail ratio, and their femoral pores average 17; hence these speci-
mens are intermediate.
Crotaphytus wishizenu. from Humboldt County, Nevada, has 18 -+
femoral pores on the average, and a body-tail ratio of .504 (average
of four specimens, agreeing closely with figures derived from Richard-
son’s (1915, p. 407) measurements, of .496 for males and .510 for
females of this species from Pyramid Lake, Nevada. The same species
from southeastern California has a body-tail ratio of .463 (average
of five specimens) and an average of 23.7 femoral pores. The mini-
mum body-tail ratio in the Nevada specimens is .500, and the maximum
in the southern California specimens measured is .492. The maximum
number of femoral pores in the northern specimens is 19, the mini-
mum in the southern examples 20.
Novres ON CNEMIDOPHORUS TIGRIS
Because of the practical impossibility of separating Cnemidophorus
tigris and Cnemidophorus stejnegert at certain points on the desert
divides and farther east in southern California, the writer believes that
the forms in question had best be placed together as subspecies. Our
series of Cnemidophorus (excepting beldingi) includes about 430
specimens from California and northern Nevada. Two critical locali-
ties, Walker Pass, Kern County, and San Gorgonio Pass, Riverside
County, are represented by large numbers of specimens. <A study of
these shows that there is both an intergradation of Cnemidophorus
tigris mundus (a new name here proposed for (. tigris undulatus
preoccupied by the C. wndulatus of Wiegmann, 1834, pt. 1, pp. 27-28)
with tigris in the Walker Pass region, as indicated by Stejneger (1893,
pp. 200-201), and of stejnegeri with tigris in Antelope Valley, Los
Angeles County, around the north base of the San Jacinto Mountains
(cf. Atsatt, 1913, pp. 89-40), and in eastern San Diego County.
The Walker Pass intergrades show a puzzling similarity to
stejnegert in the heavy spotting of the throat and, as a rule, in the
lack of smaller central gular scales. Since none of the specimens from
the Walker Pass region have as large gular scales as some stejnegeri,
and in view of their geographic position linking two closely related
subspecies, it seems better to consider them as above, and to suggest
72 University of California Publications in Zoology [Vou 17
that the color characters of stejnegert, like some factors of its environ-
ment, are intermediate between those of tigris and of mundus.
The difficulty of separating tigris and stejnegeri comes from the
fact that stejnegert on the desert borders of its range gradually takes
on the dusky suffusion of tigris, and that many examples of tigris,
even from the interior desert regions, have fully as large central gular
scales as the majority of sfejnegert.
Intergrades between stejnegert and mundus have not been recog-
nized in our material. Specimens from Fairmont, on the Mohave
Desert, Los Angeles County, and from Matilija, Ventura County, are
stejnegeri; and others from San Emigdio Plain, Kern County, and
Santa Margarita, San Luis Obispo County, have the central gular
scales smaller and are placed with mundus. We have no specimens
from the interlying area.
VARIATIONS OF EHUMECES SKILTONIANUS
Van Denburgh (1896, p. 350) has described a large, red-headed
skink from the Yosemite Valley as Eumeces gilberti. In referring to
this species later (1897, p. 149) the same writer makes the statement
that ‘‘Were it not for the different position of the light stripes of
the young and the fact that this form seemingly does not oceur in
most parts of the range of EF. skiltonianus, Ewmeces gilberti might be
regarded as a color phase of the Western Skink.’’ Cope (1900, pp.
640-644), deseribing two new ‘‘varieties’’ (amblygrammus and
brevipes) of EF. skiltonianus from California, apparently ignores the
name gilberti, but mentions a ‘“‘largest’’ specimen of FE. s. skiltonianus
“
from El Dorado, California, in which the ‘‘upper parts are entirely
”
uniform olive. The question still remains, therefore, whether
“*gilberti’’ may not be a variation of FE. skiltonianus due to age, rather
than a distinct species.
The Museum of Vertebrate Zoology has a series of eighty-one
Eumeces from the various parts of California, including thirty speci-
mens of the form ‘‘gilbertz’’ from near the type locality of the latter,
in Stanislaus, Merced, Mariposa, and Madera counties. There are also
in this series ten large specimens of Ewmeces, not to be distinguished
from ‘‘gilberti,’’ from Amador, San Joaquin, Kern, San Bernardino
and San Diego counties. All of our skinks from the coast districts
north of San Diego County are under seventy-five millimeters in body
length and have distinet stripes. Females of ‘‘gilberti’’ are smaller
-~l
oo
1916 | Camp: Subspecies of Sceloporus occidentalis
than the large red-headed males and all but one (this specimen from
the Kern River, twelve miles below Bodfish) retain some traces of the
primary longitudinal striping. Of our seventeen adult ‘‘gilberti’’
from near the type locality only four are females and these are the
smallest of the lot (maximum body length of females, ninety-two milli-
meters; of males, 107). In the coast districts the males and females
are of equal numbers and of the same size. Eight young specimens
’
from near the type locality of ‘‘gilberti’’? show as much variation in
the position of the middle pair of dorsal white lines as does skiltont-
anus from elsewhere. But a single individual from Yosemite Valley
has the white stripes separated by only the two median scale rows.
Others have from one-fourth to one-half of the second rows dark, as
do specimens at hand from other parts of California.
The upshot of the matter, then, is that all the California Eumeces
are to be considered as a single species, skilionianus, which exhibits
age and sex variations almost identical with those shown by the eastern
skink, EH. quinquelineatus. According to Cope (1900, pp. 636-637)
quinquelineatus ‘‘attains a much larger size in the more southern
states than in the northern, there going through all the stages of
coloration, and that the farther north the more is this restricted
to the primary pattern.’’ A parallel situation seems to obtain in
skiltonianus.
Transmitted October 11, 1916.
74 University of California Publications in Zoology | Vou. 17
LITERATURE CITED
AtsaTT, S. R.
1913. The reptiles of the San Jacinto area of southern California. Univ.
Calif. Publ. Zool., 12, 31-50.
Corr, E. D.
1896. On two new species of lizards from southern California. Amer.
Nat., 30, 833-836.
1900. The crocodilians, lizards and snakes of North America. Ann. Rep.
U. S. Nat. Mus., 1898, pt. 2, 151-1294, 36 pls., 347 figs. in text.
RICHARDSON, C. H.
1915. Reptiles of northwestern Nevada and adjacent territory. Proc. U.S.
Nat. Mus., 48, 403-435.
STEJNEGER, L.
1893. Annotated list of the reptiles and batrachians collected by the Death
Valley Expedition in 1891, with descriptions of new species, U.S.
Dept. Agric., Div. Ornithology and Mammalogy, N. Amer. Fauna,
7, 159-228, 394-398, 4 pls.
VAN DENBURGH, J.
1896. Description of a new lizard (Humeces gilberti) from the Sierra Ne-
vada of California. Proce. Calif. Acad. Sei. (2), 6, August 28,
1896, 350-352.
1897. The reptiles of the Pacifie Coast and Great Basin, an account of
the species known to inhabit California, and Oregon, Washington,
Idaho and Nevada. Occasional Papers Calif. Acad. Sei., 5, 236
pp., many figs. in text.
WIEGMANN, A. F. A.
1834. Herpetologia Mexicana. Pars I (Berlin: C. G. Liideritz), vi + 54,
10 pls.
ae eo A epee
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UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 8, pp. 75-114, pls. 5-12, 18 text figures March 3, 1917
OSTEOLOGICAL RELATIONSHIPS OF THREE
SPECIES OF BEAVERS
BY
F. HARVEY HOLDEN
(Contribution from the Museum of Vertebrate Zoology of the University of California)
CONTENTS
PAGE
TRISHA ODIGIY, sc eee rer SS Be RE Ce OE 76
SYEOYDO) _. cescennstes cee cAs Ae Annee CCR RS EE Be RE ep Ree 76
NL Ee ae Sener renee re 76
PACK O WIE COM ONES) c22esccscsnceccocancnesescesnecseseesesccrosteosesentesecanscasceserecseuneenaeecececarucvaseaeesore Cee
(CHERRY aE EASY ES ee 78
(CHER TIOEN ALAC OEE) ce ne
Thoracic and lumbar vertebrae
SHENG Tee RECS eS) OUT Eee ere ee ee eee eee ee eee eee
Wan CL ANRVENUe Dm 8 Cee cees cman asec ose eh kuna n ceete ssa suecccsesecadssueete retvat up casuceodseecuacecaeeencn nese
TEP SYS) ERENCES eT ae eee eee Rao Rh ee eee ea
(SOE SO cs oe 88
Clayicle . 89
1S RREPR@ TES dee aac eee seco Ree ee ee RS a EN ae oe 89
TREY GIIETS), pte ees Sete ian ee tee er Sore an eR ee ee OREN PCO EE gee RES 91
TUMNEES ge Se RS et ee fe Se eC er Ee Pee SP et een 91
(OS) THPOU ENA ATI 000, oe ee een 92
Femur
Tibia
TENANTS 5 ee en i ca g
NIELSEN EE US) so Sioa Sans ctecttesetssbeads 95
(CRI CRTAETICD ee ee a eee ae 95
Tabulation of the differences noted in the bones of the limbs and of the pectoral
Bias! ORTRIS MAGUS sect re Se eR OS EE EC So SSE See EEOC 96
RGememell MO lise revev til ON Speco cess oes ae cad cc cece aa geet ee cee ee SS 99
TL pitic@rHEy REARS: COMET cae a ee er eee 99
sonian Instig,
%
2?
%* MAR13 1917 %&
4,
“onal Muse™S
76 University of California Publications in Zoology [Vou. 17
gy
INTRODUCTION
ScoPr
The following paper contains the results of a detailed comparison
of certain extra-cranial parts of the skeletons of Castor subauratus,
Castor canadensis, and Castor fiber. This study was originally under-
taken with the first two species, to determine whether the osteological
data which could thus be obtained would confirm the systematic status
of the forms as indicated by the study of the skins and skulls alone
(see Taylor, 1916). After these observations had been recorded, a
single skeleton of Castor fiber was obtained and the condition of each
point of variance between canadensis and subauratus was noted.
These results, also, have been incorporated.
The specific characters of canadensis, as regards the skeletal parts
studied, have been taken from the three subspecies Castor canadensis
belugae, C. c. lewcodonta, and C. c. phaeus. In addition to the com-
parisons given in the text of the present paper, a tabulated statement
is added, giving the most obvious differences to be noted in the bones
of the limbs and of the pectoral and pelvie girdles.
It should be remembered that the comparisons, throughout, are
based upon a small series of skeletons, so that the examination of
further material may prove that some of the characters given are not
diagnostic. So far as it has been found feasible, all characters which
appear to be of average persistence, but which fail in one or two in-
stances in the series studied, have been noted as ‘‘tendencies.”’
MATERIAL
The major portion of the observations herein contained were made
from the following specimens now in the California Museum of Verte-
brate Zoology.
No.
Castor canadensis phaeus 210
Sex Locality
3S Hasselborg Lake, Admiralty Island,
Alaska
Castor canadensis belugae 4347 ¢ Snug Harbor, Alaska Peninsula,
Alaska
Castor canadensis leucodonta 12101 @ Alberni, Vancouver Island, B. C.
Castor subauratus 8869 ¢ San Joaquin River, near Grayson,
Stanislaus Co., California
2 San Joaquin River, near Grayson,
Stanislaus Co., California
Castor subauratus 8987
1917] Holden: Osteological Relationships of Beavers LOC
No. Sex Locality
Castor subauratus 12654 9 San Joaquin River, near Grayson,
Stanislaus Co., California
Castor subauratus 12668 og San Joaquin River, near Grayson,
Stanislaus Co., California
Castor subauratus 16385 g San Joaquin River, near Grayson,
Stanislaus Co., California
Castor fiber 19229 ? Elbe River, near Wittenberg, Ger-
many
There will also be found occasional references to other skeletons
in the Museum which as yet have not been prepared for study. These
are:
No. Sex Locality
Castor canadensis leucodonta 12107 Alberni, Vancouver Island, B. C.
Castor subauratus 8988 San Joaquin River, near Grayson,
Stanislaus Co., California
Castor subauratus 16382 2 San Joaquin River, near Grayson,
Stanislaus Co., California
Castor subauratus 16383 San Joaquin River, near Grayson,
Stanislaus Co., California
Castor subauratus 16384 San Joaquin River, near Grayson,
Stanislaus Co., California
It has been impossible up to the present time to obtain a skeleton
of Castor canadensis frondator, which form is found in parts of the
southwestern United States and barely enters California along the
Colorado River.
As far as can be determined from the skeletons, with two excep-
tions, there is but little difference in age among the specimens of
subauratus ; no. 12654 is much the oldest, and no. 16384 is a juvenile.
Among the four representatives of the subspecies of canadensis the
specimen of phaeus appears to be slightly older than the other three.
ACKNOWLEDGMENTS
It is proper to make acknowledgment here of many suggestions
kindly proffered by Professor William E. Ritter. The writer would
also extend thanks for various suggestions to Professors John C. Mer-
riam, Charles A. Kofoid, and J. Frank Daniel, as well as to Doctors
Joseph Grinnell and Walter P. Taylor, of the staff of the California
Museum of Vertebrate Zoology.
78 University of California Publications in Zoology [Vou. 17
COMPARISONS
CERVICAL VERTEBRAE
In all the cervical vertebrae there is a tendency for the laminae
and the pedicles to be heavier in Castor fiber and Castor subauratus
than in Castor canadensis belugae, C. c. leucodonta, or C. c. phaeus.
The neural canal appears to be more constant in shape throughout the
series of subauratus and in fiber than in the three subspecies of cana-
densis. There is a tendency for the articulating face of the post-
Fig. A.—Dorsal view of atlas of Castor fiber, no. 19229. X 1.
Fig. B.—Dorsal view of atlas of Castor subauratus, no. 12654, X 1.
Fig. C.—Dorsal view of atlas of Castor canadensis phaeus, no. 210. X 1.
zygapophysis of subauratus and fiber to be more convex and to face
laterad to a greater degree than it does in the subspecies of canadensis.
The average anteroposterior width of the most dorsal portion of
the atlas is greater in subauratus and fiber, the average for the three
subspecies of canadensis being 7.8 mm., for subauratus 9.8 mm., and
for fiber 8.9 mm. (see text-figs. A, B, C). In the three forms of cana-
densis the transverse processes of the atlas are thinner and constitute
a greater proportion of the entire transverse width than the same
processes of swbauratus. The odontoid process of the axis is most
highly developed both in diameter and in length in phaeus; sub-
auratus and fiber are next in size in this particular, and belugae and
leucodonta are both smaller (see text-figs. D, E, F). The width of
the anterior articulating face, at the base of the odontoid process,
1917] Holden: Osteological Relationships of Beavers 79
shows a very marked tendency to be greater in swhauratus and fiber
than in the three subspecies of canadensis (see text-figs. D, E, F) ; the
average width for the last three forms is 21.8 mm., for swbauratus
26.1 mm., and for fiber 25.7 mm. The neural canal in the axis is
Fig. D.—Cephalic view of axis of Castor fiber, no. 19229. X 1.
Fig. E.—Cephalie view of axis of Castor subauratus, no. 12654. X 1.
Fig. F.—Cephalie view of axis of Castor canadensis leucodonta, no. 12101.
x 1.
larger in subauratus than in the three subspecies of canadensis, the
canal in fiber being intermediate in size (see text-figs. D, E, F).
The neural canal in the third cervical vertebra of subawratus is
nearly semicircular in shape, while in fiber and the three subspecies
Fig. G.—Cephalic view of third cervical vertebra of Castor fiber, no. 19229.
x I
Fig. H.—Cephalic view of third vertebra of Castor subauratus, no. 8987. X 1.
Fig. I.—Cephalic view of third cervical vertebra of Castor canadensis leuco-
donta, no. 12101. X 1.
of canadensis the laminae and pedicles form a segment of a circle
which is greater than a semicircle (text-figs. G, H, 1). The vertebr-
arterial canal of swbawratus in the third cervical vertebra is almost
circular in shape, while in the three subspecies of canadensis the canal
80 University of California Publications in Zoology [Vou.17
tends to be elliptical, with the major axis extending transversely
(text-figs. G, H, 1). There appears to be a tendency in belugae and
phaeus for the vertebrarterial canals to persist throughout the entire
seven cervical vertebrae; in lewcodonta they end with the sixth; in
subauratus there are two foramina, as a rule, in the sixth cervical
vertebra, while the seventh bears one in the right transverse process
only; and in fiber, in the one specimen observed, there was one fora-
men in the left process of the seventh thoracic vertebra. From this
it will be seen that lewcodonta alone accords with the specimens of
canadensis described by Morgan (1868, p. 51) in having foramina
throughout six of the series only.
Number of differences noted between Castor subauratus and
Castor canadensis, thirteen. Fiber conforms with subawratus in nine
of these, and with canadensis in one; three remaining unassigned.
THoracic AND LuMBAR VERTEBRAE
Castor fiber and the three subspecies of Castor canadensis follow
the usual rule among rodents and the recorded descriptions of beavers
in having nineteen trunk vertebrae (Owen, 1866, pp. 364, 365; Flower,
1876, p. 50; Morgan, 1868, p. 51; Reynolds, 1897, p. 19). With the
exception of one specimen, lewcodonta, no. 12101, they consist of four-
teen thoracic and five lumbar vertebrae (Morgan, 1868, p. 19); the
individual designated, however, has thirteen thoracic and six lumbar
vertebrae, which, according to Owen (1866, p. 365), is generally the
case among rodents, and holds with but few exceptions. It is to be
noted, however, that a specimen of lewcodonta (no. 12107) has four-
teen thoracic and five lumbar vertebrae, so the possession of thirteen
thoracic vertebrae cannot be used as a character of lewcodonta.
In the series of swbauratus, without an exception, the number of
trunk vertebrae is twenty. In the nine skeletons of this species at
hand, there are four with fifteen thoracie and five lumbar vertebrae ;
four with fourteen and six; and one (no. 16382) with fourteen well-
developed ribs articulating on the fourteen thoracic vertebrae, and a
rudimentary rib on the right side of the first lumbar vertebra. It
may be worthy of note, also, that of the five males three have fifteen
thoracic vertebrae, the exceptions being nos. 16383 and 16385; and
among the four females three have fourteen thoracic vertebrae, one
of these having the variation noted above (no. 16382), and no. 8988
with fifteen thoracie vertebrae. This may show that there is a tend-
1917] Holden: Osteological Relationships of Beavers $1
eney toward sexual variation in subawratus in this regard. The proof
of this tendeney would require for demonstration a large series of
subauratus.
Beside the differences recorded above, there are others which are
seen when the thoracic and lumbar vertebrae are taken as separate
eroups. There is a marked tendency for fiber and the three subspecies
of canadensis to have larger centra in all three dimensions, in the
thoracic vertebrae, than has swbawratus (text-figs. J, K, L, and pl. 5).
This character is accompanied in the first thoracic vertebra by a
broader neural canal in subauratus and fiber than in canadensis (pl.
5, figs. 1, 2, 3, 4) ; although the height of the canal is also greatest in
subauratus, 1 fiber it is similar to that in canadensis. In the verte-
J K [E
Fig. J.—Lateral view of fourth thoracic vertebra of Castor fiber, no. 19229.
x1.
Fig. K.—Lateral view of fourth thoracic vertebra of Castor subauratus, no.
16385. X 1.
Fig. L.—Lateral view of fourth thoracic vertebra of Castor canadensis belugae,
no. 4347. X 1.
brae which follow, however, there is a marked tendency for phaeus
to have the largest neural canal (pl. 5, figs. 5, 6, 7, 8). In the eighth
thoracie vertebra and the thoracic vertebrae which follow, phaeus
possesses the largest neural canal, both in transverse and dorso-ventral
measurements.
The ratio of the width to the height of the eighth thoracic vertebra
averages 83.3 per cent in subawratus, 84.5 per cent in fiber, and 69.7
per cent in canadensis. On the thirteenth vertebra these ratios are:
subauratus, 66.4 per cent; fiber, 100.9 per cent; and canadensis, 58.6
per cent.
If we compare the height of the thoracic vertebrae from the base
of the neural spine to the most ventral portion of the centrum (ef.
§2 University of California Publications in Zoology | Vou.17
Hue, 1907, pl. 17, ‘‘S S’, hauteur totale du corps de la vertebre”’
““corps’’ does not correspond to the
BNA term corpus), we find that anteriorly in the series the greater
measurement is found in swbauratus and fiber (pl. 5, figs. 1, 2, 3, 4);
(it should be noted that Hue’s
on the first thoracic vertebra: swhauratus 16.1 mm., fiber 15.2 mm.,
and canadensis 14.3 mm. On the fourteenth thoracie vertebra, which
has a larger measurement in all three species, the opposite order holds:
canadensis 21.0 mm., subawratus 20.7 mm., and fiber 20.5 mm. Corre-
lated with this is the extreme width of the thoracic vertebrae; in the
earlier members of the series the greatest average measurement is found
in fiber, next in width is subauratus, and the narrowest is canadensis.
Later in the series, however, fiber, canadensis, subauratus, is the order
(pl. 5).
There are certain structural features which are differently located
or differ in extent in the three species. The neural spines of the first
ten thoracic vertebrae in suwbauratus and_ fiber are styliform, while
the remainder are flattened from the sides; in the three sub-
species of canadensis only the first nine are styliform, the remainder
being laterally compressed. The thoracic vertebrae posterior to the
eleventh or twelfth, in swhawratus, have an articulating surface be-
tween the anapophysis and the prezygapophysis; on the other hand,
there are no articulations between these processes in canadensis or
fiber.
On the dorsal surface of the transverse process of the first thoracie¢
vertebra of all three species of beaver so far as has been observed,
there is a small tuberculum at the place where the anapophysis joins
the metapophysis. In belugae, on the second thoracic vertebra and
on several posterior to it, this process is ear-shaped (text-fig. L) ; in
leucodonta this ear-shape appears first on the third thoracic vertebra ;
in phaeus on the fourth; in subawratus and fiber this ear-shape does
not occur on any of the first six (text-figs. J, K), but is seen first on
the seventh thoracic vertebra. It is noteworthy that these differences
show merely a tendeney in the three subspecies of canadensis for this
ear-shaped process to occur farther anteriorly in the series of thoracic
vertebrae. More material would be necessary to prove the exact loca-
tion of the first appearance of this form of the joined processes in each
subspecies of canadensis.
In subauratus the neural spines of the thoracic vertebrae, with
the exception of the first, are heavier than in the three subspecies of
canadensis (text-figs. K, L).
1917] Holden: Osteological Relationships of Beavers 83
There is a tendency for swbauratus and fiber to have ten clearly
defined pairs of transverse processes, and for the three subspecies of
canadensis to have only nine. One specimen of the latter possesses
only eight.
The neural spine of the first thoracic vertebra is shorter and less
inclined from the vertical in suwbauratus than in fiber or the three
subspecies of canadensis (pl. 5, figs. 1, 2, 3, 4). On the first thoracie
vertebra of fiber, subauratus, and phaeus there is a small notch on
the posterior margin of the vertebral arch at the base of the neural
spine, which does not appear in the other two forms. It is possible
to distinguish subauratus from phaeus or fiber by this character, as
the notch in phaeus and fiber allows the neural spines of the first two
thoracic vertebrae to come very close together along the median plane,
while in subauratus the neural spines are held apart.
There is a tendeney for the neural spine of the fourth thoracic
vertebra to be longer in subauratus and fiber than in the three sub-
species of canadensis.
In the three forms of canadensis the ear-shaped processes on the
eighth thoracic vertebra extend laterally to a greater degree than the
transverse processes, while in subawratus and fiber the opposite is
true. In subauwratus, leucodonta, and belugae, on the seventh and
eighth thoracie vertebra, the right postzygapophysis is shorter than
the left process. This is not true in phaeus or fiber, in which forms
they are of equal length.
There is a tendency for the rudimentary metapophysis to occur
on the tenth, eleventh, and even on the twelfth thoracie vertebra of
subauratus, while in belugae, fiber, and leucodonta this process is
found on the ninth and tenth thoracie vertebrae, or on the tenth
only. This process articulates with the postzygapophysis of the pre-
ceding vertebra. The exact location of this process is as follows:
subauratus, tenth and eleventh, nos. 12654, 12688, and 16385; eleventh
only, no. 8987; eleventh and twelfth, no. 8869; lewcodonta, ninth and
tenth, no. 12101; belugae and fiber, tenth only, nos. 4347, and 19229.
This process does not occur, so far as the material at hand shows,
in phaeus.
On the tenth thoracic vertebra of phaeus there is a tendency for
the postzygapophysis to bifureate and to have a dorsal as well as a
ventral articulation with the prezygapophysis of the eleventh thoracic
vertebra. :
There is a tendency for the last four thoracic vertebrae of swb-
auratus and fiber to have the articulating faces of the prezygapophyses
84 University of California Publications in Zoology [Vou. 17
and postzygapophyses inclined from the horizontal more than are
the articulating faces in the three subspecies of canadensis.
In subauratus and fiber, on the thirteenth thoracic vertebra, the
anapophyses are heavier than the same processes of the three sub-
species of canadensis. In the last-named species and fiber, however,
they are longer than in subauratus.
The average lateral extension of the metapophyses and ana-
pophyses, measured from the lateral tip of the left to the lateral tip
of the right process, and the ratios of the two widths at the thirteenth
thoracie vertebra, are as follows:
Ratio of widths
of metapophyses
Metapophyses Anapophyses to anapophyses
Castor fibers se 21.88 mm. 33.74 65%
Castor SUbaUnatIS, pecceseee eee 25.4 29.1 89
Three subspecies of Castor cana-
densis; 0.46). eee 24.2 30.5 79
C7)
YY
Fig. M.—Dorsal view of second lumbar vertebra of Castor fiber, no. 19229.
Fig. N.—Dorsal view of second lumbar yertebra of Castor subawratus, no.
16385. X 3%.
Fig. O.—Dorsal view of second lumbar vertebra of Castor canadensis belugae,
no. 4347. X %.
It will be noted from this table that the ratio of the width of the
metapophyses is nearer unity in swbawratus than m the three sub-
species of canadensis or fiber.
In the lumbar vertebrae the transverse processes of the three sub-
species of canadensis have a greater spread transversely than have
those of subauratus (text-figs. M, N, O, P, Q, R). The shape of these
processes also differs, the transverse processes in subawratus and fiber
being nearly uniform in width from their origin at the pedicles to
their extreme lateral portion; those of belugae taper near their lateral
extremity, and in leucodonta and phaeus there is a sharp point at the
very end (text-figs. M, N, O).
The mesial edges of the postzygapophyses of the lumbar vertebrae
in subauratus form an angle with each other, while in the three sub-
1917] Holden: Osteological Relationships of Beavers 85
species of canadensis these edges form one continuous are (text-figs.
M, N, 0). In fiber these edges, on the first two lumbar vertebrae,
form an angle; but on the remaining vertebrae these edges are uneven.
In the lumbar vertebrae of swhauratus and fiber there are one or two
well-developed foramina on the ventral surface of the centra, while in
the three forms of canadensis these foramina appear only on the last
or the next to the last vertebra.
In the series of swbauratus the anapophyses, which occur on the
first, second, and even the third lumbar vertebrae, have a tendency
to form an articulating surface with the metapophyses following (text-
figs. M, N, O) ; in fiber and in the three subspecies of canadensis these
processes do not articulate with each other.
Fig. P.—Caudal view of second lumbar vertebra of Castor fiber, no. 19229.
xX %.
Fig. @.—Caudal view of second lumbar vertebra of Castor subauwratus, no.
16385. X %.
Fig. R.—Caudal view of second lumbar vertebra of Castor canadensis belugae,
no. 4347. X %.
Although the prezygapoplhyses of the third and the next to the last
lumbar vertebrae extend laterally to a greater degree in subauratus
and fiber than in the three subspecies of canadensis, the postzyga-
pophyses of the same vertebrae extend laterally to a greater degree
in phaeus than they do in belugae, subauratus, or leucodonta, while
fiber exceeds even phaeus in this measurement.
The prezygapophyses of the last lambar vertebra of phaeus spread
laterally to a greater degree than do the same processes of subauratus,
which in turn exceed those of belugae and leucodonta.
Number of differences noted between Castor subauratus and Castor
canadensis, thirty-six. Castor fiber conforms with subauratus in
twenty-one of these, and with canadensis in thirteen; two are un-
assigned.
56 University of California Publications in Zoology |Vou. 17
SacRAL VERTEBRAE
The term sacral vertebrae, as used throughout this paper, includes
those vertebrae between the lumbar and caudal vertebrae which are
firmly fused together and one or more of which are joined to the ilia.
It is in this sense that the term is used by Owen (1866, p. 366),
Morgan (1868, p. 51), and Flower (1876, p. 62); and this is essen-
tially the definition given by Reynolds (1897, p. 16), although on
page 452 of the same work he says that beavers have but one sacral
vertebra.
There are five points where fusion between the sacral vertebrae
may occur: the lateral extremities of the transverse processes; the
prezygapophyses and postzygapophyses; the pedicles; the neural
spines; and the centra (pl. 6).
Phaeus, belugae, and one specimen of lewcodonta (no. 12107) have
the first four vertebrae following the lumbars fused at all five points
(pl. 6, figs. 14, 15). The second specimen of leucodonta (no. 12101)
has the first three fused at all five points, but the fourth is fused
to the third at the zygapophyses and the neural spines only (pl. 6,
fig. 16). From this it will be seen that there is a decided tendency
toward four sacral vertebrae in the three subspecies of canadensis.
In subauratus, however, three is the usual number (pl. 6, figs. 9,
10, 11, 12). In none of the specimens of the latter are the centra
of more than three vertebrae fused together (no. 16385 has only
two), although no. 12668 has a peculiar dorsal epiphysis that joins
the neural spines of the third and fourth vertebrae. The only
specimen of subauratus which does not show three and only three
sacral vertebrae is no. 8987, which has the third and fourth vertebrae
fused at the neural spines and the pedicles.
Leucodonta, no. 12107, has well-developed rudimentary ribs, such
as are to be seen in Hrethizon, joming the second sacral vertebra to
the ilia. This condition exists, but to a lesser degree, in nos. 210,
4347, 19229, and 12101, of phaeus, belugae, fiber and leucodonta, re-
spectively. Subauratus does not show these rudimentary ribs in any
of the specimens (pl. 6). From this it will be seen that, if Gegenbaur’s
definition (Flower, 1876, p. 25) for the sacral vertebrae were used,
the three subspecies of canadensis would have two sacral vertebrae
and subawratus one.
The neural spine of the three subspecies of canadensis on the first
sacral vertebra is almost perpendicular, while in subawratus and fiber
this process inclines slightly caudad.
1917] Holden: Osteological Relationships of Beavers 87
With but one exception (no. 12654), swbauratus and fiber have
thickened lateral extremities on the transverse processes of the second
and third sacral vertebrae, while the extremities of these processes in
the three subspecies of canadensis come to a thin edge (pl. 6).
Number of differences noted between Castor subauratus and Castor
canadensis, four. Castor fiber conforms with canadensis in two, and
with swbawratus in two.
CaupaL VERTEBRAE
It has been impossible to ascertain the number of vertebrae in the
caudal region of several of the specimens, since some segments have
been lost from the caudal extremity. The following specimens have
their full complement of caudal vertebrae :
Caudal Sacral
No. vertebrae vertebrae
Cas tonmfib eri ssstee st nsec 19229 25 4
Gwen leucodonta: 2s..---: ease 12107 26 4
C. subauratus (adult) ............ 12654 26 3
C. subauratus (adult) -.... --- 16385 27 3
C. subauratus (juvenile) 16384 28 3
It will be seen that the number of caudal vertebrae is not constant
in subauratus. The specimens described by Morgan (1868, p. 51),
probably Castor canadensis canadensis, and Flower (1876, p. 67),
probably the same species, had twenty-five caudal vertebrae.
The lateral extension of the postzygapophyses of the first three
caudal vertebrae is greater in subauratus than in the three subspecies
of canadensis. There is also a tendency for the prezygapophyses in
subauratus to have greater lateral extension than those in fiber or
canadensis.
Only the first four caudal vertebrae of swbauratus bear neural
spines, while in phaeus and fiber the first seven, and in lewcodonta and
belugae the first six have this process.
The postzygapophyses of the first five or six of the caudal vertebrae
project posterior of the centra in subauratus and fiber, but in the three
subspecies of canadensis these processes project beyond the centra on
only the first three.
There is a tendeney for the prezygapophyses and the postzyga-
pophyses of subauratus and fiber to be more nearly perpendicular to
the dorsoventral axis, while the three subspecies of canadensis have
these articulating faces set at an angle to this axis.
lee)
University of California Publications in Zoology {Vou.17
As a rule, the transverse processes which extend farthest laterally
are borne on the first caudal vertebra in canadensis; in subauratus on
the fourth; and in fiber on the second.
Number of differences noted between Castor subawratus and Castor
canadensis, eleven. Fiber conforms with subauratus in three; with
canadensis in four; and four are indeterminate.
Riss AND STERNUM
There is a tendency for the first six ribs in the three subspecies of
Castor canadensis and in Castor fiber to be flattened upon the trans-
verse axis to a greater degree than are the ribs of subawratus.
Owen (1866, p. 364) gives the number of ribs in Castor as fifteen
pairs. This is the number found in subauratus, but not in fiber or
in the three subspecies of canadensis, which have fourteen. There are,
however, only seven pairs of true ribs in swbawratus and the three
subspecies of canadensis; while Owen (1866, p. 366) gives the number
in Castor as eight. Morgan (1868, p. 52), who was deseribing cana-
densis, records seven pairs of true ribs.
The sternum of swbauratus and fiber is broader and thicker than
is the sternum in the three forms of canadensis.
Number of differences noted between Castor subauratus and Cas-
tor canadensis, four. Fiber conforms with subauratus in two of these
and with canadensis in two.
ScAPULA
The suprascapular border tends to be straighter in swbauratus
and fiber than in the three subspecies of canadensis (pl. 7). In the
last named forms the vertebral margin (BNA) follows the are of a
circle with its center located near the glenoid border, while in swb-
auratus and fiber the center of the are of this border is located near
the spine (pl. 7). The glenoid border is thicker in swbauratus than
in the three forms of canadensis.
The spine tends to originate at its dorsal extremity closer to the
coracoid border in fiber and in the three forms of canadensis than
it does in subauratus (pl. 7). About half-way between the acromion
and the suprascapular border there is a decided ventral twist to the
lateral portion of the spine in the three forms of canadensis, while the
spine of swhauratus and fiber is comparatively straight (pl. 7). The
1917] Holden: Osteological Relationships of Beavers 59
lateral portion of the spine in subauratus and fiber has a projecting
edge on its glenoid side, which extends from the acromion about two-
thirds of the way to the suprascapular border; in the three subspecies
of canadensis this projecting edge is shorter, ending where the spine
twists ventrad.
The angle formed by the glenoid border and the coracoid border,
if produced, is greater in subauratus than in fiber or in the three
forms of canadensis, or, it might be said, these borders approach each
other more rapidly in swbauratus. Fiber and the three forms of cana-
densis have a more nearly sharp-cornered superior angle than has
subauratus (pl. 7). The coracoid process of subauratus and _ fiber
has a much sharper point than it has in the other three forms, but
it is heavier in canadensis and fiber. The ventral extremity of the
acromion tends to be more nearly styliform and narrower in the three
subspecies of canadensis than in subauratus or in fiber.
Number of differences noted between Castor subauratus and Cas-
tor canadensis, twelve. Fiber conforms with subawratus in seven;
with canadensis in four; and one is indeterminate.
CLAVICLE
The shaft of the clavicle in Castor subauratus, if projected upon
the dorsoventral and cephalocaudal planes, would describe ares, the
center on the first of which would be located mediodorsad, and that
of the second, laterocephalad (pl. 8, figs. 30 to 37). In the three forms
of Castor canadensis and in Castor fiber the shaft is straight. On the
costal surface, near the dorsal edge and about one centimeter from
the acromial extremity, there is a small fossa, evidently for the attach-
ment of muscle; this is deeper in subauratus and fiber than in the
other forms.
Number of differences noted between Castor subauratus and Castor
canadensis, three. Fiber conforms with suwbauratus in one, and with
canadensis in two.
Humerus
The base of the epiphysis of the greater tuberosity is broader in its
anteroposterior dimension in Castor fiber and in the three subspecies
of Castor canadensis than in Castor subauratus. The bicipital groove
is longer and deeper in fiber and in the three forms of canadensis
than it is in subauratus (pl. 9). The olecranon fossa is more clearly
90 University of Califorma Publications in Zoology [Vou.17
defined in the three subspecies of canadensis than in subawratus or
fiber. The proximal surface of the deltoid ridge has a tendency to
be flattened transversely in leucodonta and phaeus, but not in sub-
auratus, fiber, or belugae. The proximal origin of the deltoid ridge
is more distally located in the three forms of canadensis than in sub-
auratus or fiber (pl. 9). There is a marked difference in the form
of the internal condyle; in fiber and in the three forms of canadensis
it is greater in its longitudinal measurement than in swbauratus, and
in subauratus its transverse dimension is greater than in fiber or in
the three subspecies of canadensis.
The supinator ridge is heavier and more inclined to have an an-
terolateral ridge in subauratus and fiber than in the three subspecies
of canadensis (pl. 9). In subauratus and fiber the supinator ridge
forms an even curve, while in the three subspecies of canadensis there
is an easily discerned angle about half-way between the shaft and the
external condyle.
There is a small tuberosity on the lateral anterosuperior surface of
the trochlea in subauratus which does not exist in any of the other
four forms (pl. 9, figs. 38, 39, 40, 41). The capitulum has a tendency
to be heavier in belugae and phaeus than in subauratus, fiber, or
leucodonta.
In subauratus and fiber the anterior distal articulating surfaces
are wider, in comparison with the greatest width of the distal ex-
tremity of the humerus, than is the case with the three subspecies of
canadensis. This is shown in the following table:
Width art. Width dist. Ratio,
surfaces extremity per cent
Subspecies of Castor canadensis (three
SP@CIMeENS))| sess sescsseccsccewecee seen seeeeee es 18.9 mm. 32.3 mm. 58.5
Castor fiber (one specimen) wns PAULS) 31.0 67.0
Castor subauratus (five specimens) ... 19.2 30.9 62.1
The distal wall of the small ellipsoidal fossa on the posterior
surface of the internal condyle is heavier in fiber and in the three
subspecies of canadensis than it is in subauratus.
The supinator ridge extends laterally farther from the distal
articulating surfaces in canadensis than in subauratus or fiber (pl. 9).
Number of differences noted between Castor canadensis and Castor
subauratus, sixteen. Fiber conforms with swbhauratus in nine and
with canadensis nm seven.
1917] Holden: Osteological Relationships of Beavers 91
Rapius
The plane of the proximal articulating surface of the radius makes
an angle of about ninety degrees with the shaft of the radius in the
three subspecies of Castor canadensis, and in Castor fiber, while in
Castor subauratus the superior margin of the proximal epiphysis is
tipped distally, so that the plane of this articulating surface makes a
more acute angle with the axis of the radius (pl. 10). The greatest
diameter of the neck of the radius is less in the three subspecies of
canadensis than in subauratus or fiber: canadensis averages 6.10 mm. ;
subauratus 7.94; fiber 7.65. On account of this greater constriction
at the neck of the radius, the head rises more abruptly from the shaft
in the three subspecies of canadensis. The most lateral portion of the
head bears a small tubercle in the three forms of canadensis which, in
subauratus and fiber, is not so well developed (pl. 10). Proximad
and mediad to the bicipital tuberosity is a well-marked fossa which is
developed to a greater degree in subauratus than in fiber or in the
three subspecies of canadensis. In the three forms of canadensis the
sigmoid cavity is deeper than it is in subauwratus or fiber.
Number of differences noted between Castor canadensis and Castor
subauratus, six. Castor fiber conforms with subauratus in four of these
and with canadensis in two.
ULNA
On the external face of the olecranon, near the posterior edge,
there is a well-defined crest in the three subspecies of Castor cana-
densis. In Castor subauratus and Castor fiber this is not developed
to such a degree, but is seen as a line rather than as a crest (pl. 10).
The anterior edge of the olecranon is thinner in the three sub-
species of canadensis than in subawratus or fiber. This is in part
due to the increased depth of the fossa located on the internal surface
of the ulna below and proximad to the greater sigmoid notch. This
fossa, in the three forms of canadensis, lies close to the superior edge
of the olecranon, while in subauratus it lies farther posteriorly from
this edge. The mesial edge of the coronoid process is thicker in sub-
auratus and fiber than in the three forms of canadensis; in the latter
this edge is sharp. The lateral edge of the lesser sigmoid cavity is
also thicker in the case of subauratus and fiber than in the three sub-
species of canadensis. Laterad to the lesser sigmoid cavity is a tuber-
culum which is less developed in subauratus and fiber than in the
forms of canadensis.
t
92 University of California Publications in Zoology [ Vou. 17
The under surface of the ulna from the point of the olecranon
to a point beneath the greater sigmoid cavity has, in the three sub-
species of canadensis, the appearance of having been roughly modeled
with a smooth, flat tool. This gives several smooth, flat surfaces, with
more or less well-defined edges between them. In subawratus and
fiber this surface is rounded, having no flat surfaces or well-defined
edges.
Immediately below the interosseous crest is a fossa which ends
rather abruptly proximad in the three forms of canadensis and in
fiber, while in subauratus it comes gradually to the surface (pl. 10).
The minimum diameter of the shaft, which occurs near the distal
extremity, is less in subauratus than in canadensis or fiber: subauratus
measures 4.60 mm.; canadensis 5.17; fiber 6.50. In the three sub-
species of canadensis the greatest diameter of the distal epiphysis
is less in comparison with the total length of the epiphysis than in
subauratus or fiber: the ratio of the diameter to length in canadensis
is 76.1 per cent; subauratus 84.8; fiber 89.3 (pl. 10).
Number of differences noted between Castor canadensis and Castor
subauratus, ten. Fiber conforms with subawratus in six, with cana-
densis in two, and two of the differences are unassigned.
Os INNOMINATUM
There is a tendency for the iliae portion of the acetabulum to be
deeper and longer anteriorly in Castor subawratus than in the three
forms of canadensis. Those portions of the ischium and pubis which
form the base of the acetabulum are thicker in subauratus and fiber
than in the three forms of canadensis. On the posterior dorsal edge
of the acetabulum the lunate surfaces project farther ventrad in
subauratus than in the three subspecies of canadensis, or in fiber
(pl. 11).
Ventral to the acetabulum is a square angle in fiber, subauratus,
and phaeus, which does not oeeur in leucodonta or belugae.
Number of differences noted between Castor subauratus and Castor
canadensis, four. Fiber conforms with canadensis in two, and with
subauratus im one.
FEMUR
In Castor subauratus the depression in the head of the femur is
deeper than in the three subspecies of Castor canadensis or in Castor
fiber. The diameter of this depression in subauratus and fiber is
1917 | Holden: Osteological Relationships of Beavers 93
greater than in the forms of canadensis. The epiphysis of the great
trochanter forms with the shaft an angle that, in subauratus and
fiber, is nearer 90 degrees than it is in any of the three subspecies
of canadensis (pl. 12). In the three last-mentioned forms and in
fiber the intertrochanteric line is more clearly defined than in sub-
auratus. The lesser trochanter in subauratus and fiber has a flattish
epiphysis which faces proximad and mesiad, while in the three forms
of canadensis the lesser trochanter ends in a moderately sharp point
(pl. 12). Directly anterior to the lesser trochanter, on the mesial
side of the shaft of the femur, is a fossa which is deeper in subauratus,
fiber, and belugae than in leucodonta or phaeus. The third trochanter
(see Owen, 1866, p. 381, for name) in the three forms of canadensis
is less blunt at its lateral extremity than in swbauratus or fiber (pl. 12).
Number of differences noted between Castor subauratus and Cas-
tor canadensis, eight. Castor fiber conforms with subawratus in six,
and with canadensis in two.
TIBIA
The tibia is longer in comparison with the length of the femur in
the three subspecies of Castor canadensis than it is in Castor sub-
auratus or Castor fiber. The difference between subauratus and
phaeus, however, is slight, as will be seen from the following table:
Ratio,
Femur Tibia per cent
(Ch Gy TOLNGNE TS eee eee 106.6 mm. 131.6 mm. 81.1
C. e. belugae and leucodonta ............ 103.9 132.8 78.2
C. subauratus (average of five) =) LOS:7 129.4 81.7
(OL, LOVEI oo eee eee ere rere Serer ee ne eee ae 108.5 132.9 81.6
Anterior to the anterior condyloid fossa there is a tuberosity
which is much sharper in the three forms of canadensis than in swb-
auratus or fiber. In all five forms of beaver the posterior face is
deeply concave; but in swbauratus and fiber the condyloid fossa ends
much less abruptly proximad than it does in the three forms of cana-
densis.
Subauratus and fiber have heavier interosseous crests than have
the three subspecies of canadensis. The malleolar groove is also wider
in subauratus and fiber than in the three forms of canadensis, but is
deeper and longer in the three last-mentioned forms and in fiber. The
posterior margin, or rim, of the malleolar groove is thinner in the
three subspecies of canadensis than in subauratus or fiber.
94 University of California Publications in Zoology [Vou. 17
At the distal extremity of the posterior face there is a process
which has a much sharper point in subauratus than in the three sub-
species of canadensis or in fiber; but this process is smaller in sub-
auratus and fiber.
Number of differences noted between Castor subauratus and Cas-
tor canadensis, ten. Castor fiber conforms with subauratus in seven
and with canadensis in three.
FiBuna
The angle which the hamular process (see Morgan, 1868, p. 54, for
name) makes with the shaft of the fibula approximates, in Castor sub-
auratus and Castor fiber, ninety degrees, and is decidedly less than
ninety degrees in the three subspecies of Castor canadensis (see pl. 8,
figs. 25 to 29).
On the proximal third of the fibula the following differences occur :
in the three subspecies of canadensis and in fiber the lateral face has
a decided tendency to be concave, while in subauwratus there is a tend-
ency toward convexity. In the three forms of canadensis and in fiber
there is a well-defined posterior face which joins the medial face in
a sharp-edged medial crest, the medial face being slightly coneave.
In subauratus the medial and posterior faces are poorly defined and
the medial crest is but a faintly indicated line (pl. 8, figs. 25 to 29).
Proximad, and directly anterior to the face for the articulation of
the tibia, the fibula in swhawratus and fiber presents a small, flat face,
while in the three subspecies of canadensis there is a sharp crest at
this point.
The distal epiphysis has proximally pointing projections which are
claw-shaped in the three forms of canadensis but are less developed
in subauratus or fiber.
On the proximal posterolateral margin of the hamular process in
canadensis is a crest which does not occur in subauratus or fiber.
On the posterior face of the distal epiphysis is a suleus which is
wider in the three forms of canadensis and in fiber than in subawratus.
The medial face of the external malleolus tends to be convex in cana-
densis, while in subauratus and fiber this articulating face presents
the appearance of having been ground and is concave.
Number of differences noted between Castor subauratus and Cas-
tor canadensis, eight. Castor fiber conforms with subauratus in five
and with canadensis in three.
1917] Holden: Osteological Relationships of Beavers 95
ASTRAGALUS
On the medial side of the body of the astragalus is a tuberosity
which is developed to a greater degree in the three subspecies of
Castor canadensis than in Castor subauratus or Castor fiber. The
groove on the medioposterior edge extends farther laterad in the three
forms of canadensis than it does in subauratus. The lateral border
of the head of the astragalus is twisted upward more in subauratus
than in the three subspecies of canadensis or fiber. The median artic-
ulating face for the caleaneum is broader posteriorly in belugae, fiber,
and phaeus than it is in leucodonta or subauratus.
Number of differences noted between Castor subauratus and Castor
canadensis, four. Castor fiber conforms with swbauratus in two, and
with canadensis in two.
CaLCANEUM
In the three subspecies of Castor canadensis there is a well-defined
medial tuberosity which is lacking in Castor subawratus and Castor
fiber. In the three forms of canadensis and in fiber the median and
anterior articulating faces make an angle anteriorly of about forty-
five degrees with the axis of the caleaneum, while in subawratus they
make an angle of about thirty degrees. There is a decided tendency
for the sustentaculum of swbauratus to be more rounded mediad than
it is in the three subspecies of canadensis.
The plantar surface of the caleaneum in the three forms of cana-
densis and in fiber is wider and more uniform in width for its entire
length than it is in swbauratus, where there is a tendency for it to
have a constriction directly below the sustentaculum.
On the distal margin of the lateral face of the body is a process
which is more highly developed in canadensis than in subauratus or
fiber. As viewed from the plantar surface, the body of the caleaneum
is less curved in subauratus and fiber than it is In canadensis.
Number of differences noted between Castor subauratus and Castor
canadensis, seven. Castor fiber conforms with suwbauratus in three,
and with canadensis in three.
Total number of differences noted between Castor subauratus and
Castor canadensis, 156. Castor fiber conforms with subawratus in
eighty-nine (58 per cent); with canadensis in fifty-three (33 per
cent) ; and fourteen (9 per cent) are not clearly subject to assignment.
[ Vou. 17
Zoology
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University of California Publicat
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97
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[ Vou. 17
Zoology
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1917] Holden: Osteological Relationships of Beavers 99
GENERAL OBSERVATIONS
It has not been possible to formulate any one general statement
covering the differences between the three species of Castor. The fact
that swbauratus has one more thoracic-lumbar vertebra might signify
that, in this character at least, swbauratus is the most primitive form.
The three subspecies of canadensis are more closely related, inter
se, than is subauratus or fiber to any one of them. Also it is to be seen
that swbawratus and fiber are more closely related than fiber and cana-
densis. The differences, however, between subauratus and fiber are so
ereat as to preclude consideration of these two forms as conspecific.
The relative similarity, however, brings forward many interesting
questions, to which no attempt is here made to give answers.
Transmitted August 4, 1916.
LITERATURE CITED
Barker, L. F.
1907. Anatomical terminology with special reference to the [BNA]. Phila-
delphia, P. Blakiston’s Sons & Co., ix + 103 + 16, 5 diagrams.
BLAINVILLE, H. M. D. DE
1839-1864. Ostéographie, ou description iconographique comparée du squel-
ette et du systeme dentaire des cing classes d’animaux vertébrés.
Paris, J. B. Bailliére et fils; New York, Braillére Brothers; ete.,
etc. Atlas, tome 4, 93 pls.
FLoweErR, W. H.
1876. An introduction to the osteology of the mammalia. London, Macmillan
& Co., xi + 344, 126 figs. in text.
Hug, E.
1907. Musée ostéologique: étude de la faune quaternaire. Ostéométrie des
mammiféres. Paris, Schleicher Fréres, 2 vols., xix + 50, 186 pls.,
2187 figs.
Morean, L. H.
1868. The American beaver and his works. Philadelphia, J. B. Lippincott
& Co., xv + 330, 1 map, 23 pls., 26 figs. in text.
Owen, R.
1866. Anatomy of vertebrates. London, Longmans, Green & Co., II, viii +
592, 3 tables, 406 figs. in text.
REYNOLDS, S. H.
1897. The vertebrate skeleton, in Cambridge Nat. Sci. Manuals. Cambridge,
University Press, xvi +559, 110 figs. in text.
TayLor, W. P.
1916. The status of the beavers of western North America, with a consider-
ation of the factors in their speciation. Univ. Calif. Publ. Zool.,
12, 413-495, 14 figs. in text.
PLATE 5
Cephalic view of the first (upper row) and the thirteenth (lower row) thoracic
vertebrae; about nine-tenths natural size.
Figs. 1 and 5. No. 19229, Castor fiber.
Figs. 2 and 6. No. 210, Castor canadensis phaeus.
Figs. 3 and 7. No. 4347, Castor canadensis belugae.
Figs. 4 and 8. No. 16385, Castor subauratus.
Note that in swbauratus and fiber the first thoracic vertebra, figures 1 and 4,
has a broader neural canal, and that in subauratus, figure 4, the height is greater ;
that the ‘‘hauteur totale du corps de la vertébre’’ in the first thoracic vertebra
is greater in subawratus and fiber, and that on the thirteenth canadensis shows
this measurement to be greatest; that there is a tendency for fiber and canadensis
to have larger centra.
[100]
Wee Perea:
a ae ih 7
A 4 te,
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PLATE 6
Ventral view of the vertebrae from the sacral region; about one-half natural
size.
Fig. 9. No. 8987, Castor subauratus.
Fig. 10. No. 12654, Castor subauratus
Fig. 11. No. 12668, Castor subauratus.
Fig. 12. No. 16385, Castor subauratus.
Fig. 13. No. 19229, Castor fiber.
Fig. 14. No. 210, Castor canadensis phaeus.
Fig. 15. No. 4347, Castor canadensis belugae.
Fig. 16. No. 12101, Castor canadensis leucodonta.
Note that the centra of the first four vertebrae of fiber, belugae, and phaeus
are fused, while in subauratus and leucodonta only three of the vertebrae are so
joined (no. 16385 having only two). The lateral portions of the transverse
processes of the third and fourth vertebrae can be seen to be thinner in cana-
densis (and one specimen, no. 12654, of swbawratus) than in fiber or subauratus.
The presence of the second rudimentary rib in fiber, and the tendeney for cana-
densis to have a second rib to the ilium, can also be seen.
[102]
PLATE 7
Lateral view of the left scapula; about one-half natural size.
Fig. 17. No. 8987, Castor subauratus.
Fig. 18. No. 12654, Castor subauratus.
Fig. 19. No. 12668, Castor subauratus.
Fig. 20. No. 16385, Castor subauratus.
Fig. 21. No. 19229, Castor fiber.
Fig. 22. No. 210, Castor canadensis phacus.
Fig. 23. No. 43847, Castor canadensis belugae.
Fig. 24. No. 12101, Castor canadensis lewcodonta.
Note that the suprascapular border is more nearly straight in fiber and sub-
auratus than in canadensis; that the spine tends to originate at its dorsal ex-
tremity closer to the coracoid border in fiber and canadensis; the lateral twist of
the spine in canadensis and the comparatively straight spine in fiber and sub-
auratus; that the angle of the glenoid and coracoid borders, if produced, is
greatest in subawratus; that the superior angle is in canadensis more angular.
[104]
TD)
PLATE 8
Costal surface of the left clavicle and the tibial surface of the left fibula;
about two-thirds natural size.
Figs. 25 and 35. No. 210, Castor canadensis phaeus.
Figs. 26 and 37. No. 12101, Castor canadensis leucodonta.
Figs. 27 and 34. No. 19229, Castor fiber.
Fig. 28. No. 8869, Castor subauratus.
Figs. 29 and 33. No. 16385, Castor subauratus.
Fig. 30. No. 8987, Castor subauratus.
Fig. 31. No. 12654, Castor subauratus.
Fig. 32. No. 12668, Castor subauratus.
Fig. 36. No. 4347, Castor canadensis belugae.
Note that the angle which the hamular process makes with the shaft of the
fibula is greater in subauratus and fiber than in canadensis; that the shafts of
the clavicles of subauratus are curved, in canadensis and fiber they are straight;
that the fossa near the dorsal edge is deeper in fiber and subauwratus than in cana-
densis.
[106]
PLATE 9
Anterior surface of left humerus; about one-half natural size.
Fig. 38. No. 8987, Castor subauratus.
Fig. 39. No. 12654, Castor subauratus.
Fig. 40. No. 12668, Castor subauratus.
Fig. 41. No. 16385, Castor subauratus.
Fig. 42. No. 19229, Castor fiber.
Fig. 48. No. 210, Castor canadensis phaeus.
Fig. 44. No. 4347, Castor canadensis belugae.
Fig. 45. No. 12101, Castor canadensis leucodonta.
Note that the bicipital groove is deeper and larger in canadensis and fiber
than in subauratus. The proximal origin of the deltoid ridge is more distally
located in canadensis than in subauratus or fiber; the longitudinal measurement
of the internal condyle is greater in fiber and canadensis; the supinator ridge
forms an even curve in fiber and subauratus, but an angle in canadensis.
[108]
Aa
on
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PLATE 10
Lateral surface of the left ulna and radius; about two-fifths natural size.
Figs. 46 and 47. No. 8987, Castor subauwratus.
Figs. 48 and 49. No. 12654, Castor subauratus.
Figs. 50 and 51. No. 12668, Castor subauratus.
Figs. 52 and 53. No. 16385, Castor subauratus.
Figs. 54 and 55. No. 19229, Castor fiber.
Figs. 56 and 57. No. 210, Castor canadensis phaeus.
Figs. 58 and 59. No. 4347, Castor canadensis belugae.
Figs. 60 and 61. No. 12101, Castor canadensis leucodonta.
Note that the plane of the proximal articulating face of the radius is set at
an angle of about 90 degrees with the shaft in fiber and canadensis, but is tilted
in subauratus; the greatest diameter of the neck of the radius is less in cana-
densis; the head of the radius rises more abruptly in canadensis. In swbauratus
the fossa below the interosseous crest is shallower, at its proximal end, than in
canadensis and fiber.
[110]
PLATE 11
View of left os innominatum, showing the acetabulum; about three-fifths
natural size.
Fig. 62. No. 16385, Castor subauratus.
Fig. 63. No. 43847, Castor canadensis belugae.
Fig. 64. No. 19229, Castor fiber.
Note that there is a square angle ventral to the acetabulum in swbauratus and
fiber, but not in belugae.
[112]
#
View of posterior surface of
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig
g.
Fig.
65.
66.
No.
No.
No.
No.
No.
No.
No.
No.
8987,
12654,
12668,
16385,
19229,
210,
4347,
12101,
Castor
Castor
Castor
Castor
Castor
Castor
Castor
Castor
PLATE 12
the left femur; about two-fifths natural size.
subauratus.
subauratus.
subauratus.
subauratus.
fiber.
canadensis phaeus.
canadensis belugae.
canadensis leucodonta.
Note that the epiphysis of the great trochanter forms an angle nearer ninety
degrees with the shaft, in subauratus and fiber than in canadensis; that in fiber
and canadensis the intertrochanteric line is more clearly defined than in sub-
auratus ; that the lesser trochanter of fiber and subauwratus has an epiphysis, but
that it does not occur in the forms of canadensis; that the third trochanter of
canadensis is sharper than that of fiber or subauratus.
[114]
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ZOOLOGY
Vol. 17, No. 9, pp. 115-125, 3 text figures February 3, 1917
NOTES ON THE SYSTEMATIC STATUS OF THE
TOADS AND FROGS OF CALIFORNIA
BY
CHARLES LEWIS CAMP
(Contribution from the Museum of Vertebrate Zoology of the University of California)
The large series of amphibians contained in the collection of the
Museum of Vertebrate Zoology at the University of California has
been at the writer’s disposal for study and has furnished opportunity
for looking into the status of certain California toads and frogs.
Through the courtesy of Dr. Leonhard Stejneger, Head Curator in
the Department of Biology of the United States National Museum,
the writer was accorded the privilege of examining specimens in that
museum, where the types of nearly all of the western species of frogs
and toads are preserved.
As a result of this study it is found necessary to consider the two
long recognized forms, Bufo halophilus and Bufo boreas, as subspecies
of one species, and to put Bufo columbiensis under the synonymy of
Bufo boreas boreas. Two new races of that confusing species, Rana
boylivi, are described, and two other western frogs, Rana draytonw and
Rana aurora, are placed together as subspecies of aurora because of
the recognition of intergrades from the coast region of northern Cali-
fornia. Notes are also given on the occurrence in California of Bu/jo
boreas nelsoni and the two subspecies of Rana pretiosa.
Toaps
Examination of about 370 toads of the several forms closely alhed
to Bufo boreas, from Alaska, Vancouver Island, eastern Washington,
western Oregon, northern Nevada, and California, has convinced the
writer that most of the characters that have been employed by various
authors, including Baird and Girard (1854a, pp. 174-175, 378-880,
18546, p. 301), Girard (1858, pp. 74-80, atlas, pl. 5, figs. 4-9, pl. 6,
116 University of California Publications in Zoology [Vou. 17
figs. 4-9), Boulenger (1882, pp. 284-285, 295-296), Cope (1889, pp.
267-271), Stejneger (1893, pp. 220-221), and Dickerson (1906, pp.
44, 113-116), to separate the so-called species boreas and halophilus
(ineluding columbiensis) cannot, in this variable group, be considered
diagnostic. It would appear that some of the characters originally
used to distinguish boreas from columbiensis are of sexual rather than
specific difference. The male of boreas has smoother skin, longer fore-
arm and hind limb, and perhaps longer first finger than the female.
Roundness of snout as contrasted with the truneated condition con-
sidered as distinguishing halophilus is probably an age character, as
also is the degree of adherence of the skin on top of the head to the
skull. Other criteria, such as width of head, distance between eyes,
and size of parotoid, seem to be individual differences. The “‘types”’
of columbiensis in the United States National Museum are all young
specimens which I am unable to separate from boreas. Therefore,
unless a hitherto unnoted difference exists between the toads of Puget
Sound and those of the Columbia River, the name coluwmbiensis must
be dropped.
The only constant characters distinguishing borcas from halophilus
in the specimens at hand are those of increase of pigmentation of the
more northern form and decrease of webbing (best indicated by spread
of hind foot) in the southern. On the basis of these two characters
one, Bufo boreas boreas,
inhabiting Washington, Oregon, northern Nevada, and northern Cali-
two subspecies of boreas may be separated
fornia as far south as Eureka, Humboldt County, Sisson, Siskiyou
County, and Mono County; the other, a southern subspecies, Bufo
boreas halophilus, oceurring from southernmost San Diego County
north to the Gualala River, Sonoma County, to Oroville, Butte County,
and to the vicinity of Independence, Inyo County. Specimens from
localities in the intervening area show that intergradation between
boreas and halophilus takes place.
The above two subspecies are not easily confused if both the type
of coloration and the ratio of spread of hind foot to total length are
taken into account, and the followimg key may be used to separate
them.
1. Back either dark and unspotted or blotched and speckled between the blotches
with small, dark dots on the light background; spread of hind foot from
end of first to tip of fifth toe usually more than 36 per cent of the total
body-length; maximum body-length 120 millimeters —_.. Bufo boreas boreas.
1’. Back usually spotted or blotched (rarely with dark speckles between the
blotches on the light background) and seldom wholly dark; spread of hind
foot usually less than 36 per cent of total body-length; maximum body-
Ten othe) Ol ernie 61: eerste eee tee nena aera Bufo boreas halophilus.
1917] Camp: Notes on the Systematic Status of Toads and Frogs 117
The two toads from Owens Valley, Inyo County, California, re-
ferred by Stejneger (18938, pp. 220-221) to Bufo boreas nelsoni,
appear to be halophilus. The rounded snout of these is paralleled
in many half-grown halophilus in a series at hand from southern
California.
Frogs
Examination of about 450 specimens of frogs, mostly contained
in the Museum of Vertebrate Zoology, warrants an attempt to formu-
late a key to the Pacific Coast species—Rana pretiosa, Rana aurora,
and Rana boyli, and their subspecies. It has been thought necessary
to redefine Rana boyli and to describe two new races of this species,
one from the high Sierra Nevada and the other from the mountains of
southern California.
Rana boylii boylii Baird
California Yellow-legged Frog
Type locality—California (Baird, 1856, p. 62) ; subsequently in-
dicated as El Dorado, California (Cope, 1889, p. 447).
Synonyms—Rana pachyderma Cope (1884, pp. 25-27), (types
from MeCloud River |= Baird, Shasta County], California, and Ash-
land, Oregon) ; Rana pretiosa (Yarrow and Henshaw, 1878, p. 1632),
part |Kern River]; [?] Rana temporaria pretiosa (Cope, 1889, p. 434),
part [Santa Barbara].
Description.—V omerine teeth rudimentary, on two oblique ridges
nearly meeting between and behind the nares (see fig. 3) ; tympanic
region not darker than rest of head; upper labial ridge mottled or
colored like rest of body and not distinctly lighter than rest of head;
red never present in coloration (except in diseased individuals). The
above characters are diagnostic of all three subspecies of boylii. Those
characters which pertain to this subspecies alone are: hind leg long,
inside angle of bent tarsus reaching at least to nares and usually beyond
when leg is advanced along body; tibia elongate, reaching usually be-
yond anus when flexed and held at right angles to axis of body ; fourth
toe on reflexed hind foot never reaching beyond end of knee and often
not quite to fold of skin below knee; head broad and pointed when
viewed from above, its width two and one-third to two and two-thirds
times in body-length; skin on back, legs and tympanum, thick and
rough with minute brownish spines; color dorsally varying from nearly
118 University of California Publications in Zoology [Vou. 17
uniform black to ight gray, greenish or brownish, with darker mark-
ings, if present, usually indistinct; there is always a patch of lighter
color on top of head between nares and eyes, and behind this a darker
area crossing posterior half of each eyelid and merging insensibly be-
hind into the general dorsal coloration. Boylii is the smallest of the
three subspecies, reaching a maximum body-length of only about 66
millimeters.
Material—One hundred and seventy-seven specimens of R. b.
boylw are contained in the Museum of Vertebrate Zoology, from the
following localities in California: Cuddeback, Humboldt County;
eight miles east of South Yolla Bolly Mountain, Trinity County ;
vicinity of Covelo, vicinity of Sherwood, near Mount Sanhedrin, and
Gualala, Mendocino County ; Gualala River, Freestone, and eight miles
west of Cazadero, Sonoma County; Fairfax, San Anselmo, Lagunitas
Creek, and Muir Woods, Marin County; Mill Creek near Tehama,
Tehama County ; Chambers Ravine near Oroville, Butte County ; Wins-
low near Fruto, Glenn County ; near Vacaville, Solano County ; vicinity
of Mount Diablo, Contra Costa County; Berkeley, Alameda County ;
Corral Hollow, San Joaquin County ; Sweeney’s ranch, 22 miles south
of Los Banos, Mereed County; Fyffe (3600 feet altitude), El Dorado
County; Pleasant Valley (600 feet), vicinity of Coulterville (3000-
3200 feet), and near Feliciana Mountain (3800 feet), Mariposa
County; Farrington’s ranch (6800 feet), near Williams Butte, Mono
County; and Kern River near Bodfish, and Fay Creek (4100 feet),
six miles north of Weldon, Kern County.
Distribution Except for a small area in southwestern Oregon,
the range of Rana boylw boylii seems to lie entirely within the state
of California. It includes the northwestern part of the state, east to
the MeCloud River, Shasta County, and to the western foothills of the
Sierra Nevada below 4100 feet altitude. The form oceurs also at
Mono Lake; in the Sierras it has been taken south to the vieimity of
Walker Pass, Kern County; but where it meets the range of R. b.
muscosa along the coast is not known. Specimens from the vicinity
of Walker Pass show a more contrasted pattern of coloration, possibly
indicating approach to the southern subspecies, deseribed next below.
Rana boylii muscosa, new subspecies
Sierra Madre Yellow-legged Frog
Type.—Female, adult; no. 771, Mus. Vert. Zool.; Arroyo Seco
Canon, at about 1300 feet altitude, near Pasadena, California; August
3, 1903; colleeted by J. Grinnell.
1917] Camp: Notes on the Systematic Status of Toads and Frogs 119
Diagnosis—Like Rana boyli boyli, but attaining much larger
size, and (except In young) with no light patch in front of dark areas
across upper eyelids. Dorsal ground color usually lighter than in
R. b. boylv, light yellow to brown, contrasting with the darker moss-
like patches on the back. Tips of toes more expanded than in boylit.
Description —V omerine teeth on two oblique ridges between nares ;
head pointed in outline as viewed from above, broad, its width entering
body-length two and two-thirds times; hind limbs long, posterior side
of bent tarsus reaching forward to snout; fourth toe on hind foot
reaching forward not quite to knee-fold; dorso-lateral fold indistinet,
not pitted anteriorly ; tympanum and area surrounding it very rough,
beset with small tubercles; web of hind foot extending nearly to tips
of toes ; outer and inner metatarsal tubercles distinet ; plantar tubercles
very large; tips of toes expanded, dise-shaped; distal end of flexed
tibia held at right angles to body reaching anus; color above dark
yellow, with a reticulated pattern of moss-hke dark patches; beneath,
yellow, spotted with dusky on throat and chest; upper lip below eye,
mottled.
Material—F ifty-one specimens in the Museum of Vertebrate Zool-
ogy, from the following localities in southern California: San Jacinto
Mountains: Keen’s Camp (nos. 3804, 3805) ; Strawberry Valley, 6000
feet altitude (mos. 534, 550, 584) ; Fuller’s Mill, 5600-5800 feet (nos.
278-287, 314) ; Sehain’s Ranch, 4900 feet (no. 353); Cabezon, 2000
feet (nos. 151, 177); and Snow Creek, near Whitewater, 2500 feet
(nos. 79, 194-197); San Bernardino Mountains: Santa Ana River,
5500 feet (nos. 714, 715) ; Fish Creek, 6500 feet (nos. 772-774) ; and
Barton Creek, 6000 feet (mo. 4389); and San Gabriel Mountains:
Canons near Sierra Madre, 1200-3000 feet (nos. 4874-4377, 4388, 4855,
4856, 4868) ; West Fork San Gabriel River, 3000 feet (nos. 4378-4385) ;
Arroyo Seco Canon near Pasadena (nos. 770, 771); and Little Rock
Creek Canon, 4700 feet (nos. 4390, 4391).
Variations —The dorsal color pattern in some specimens, particu-
larly those from the San Gabriel Mountains, is different from any-
thing found in R. b. boylit. The ground color is lighter in these speei-
mens than in either of the two northern subspecies, and is thickly
marked with the lichen-like dark patches from which the name is
derived. In other specimens, especially those from the San Jacinto
Mountains, the color is darker, becoming uniformly dark brown in a
single individual from Strawberry Valley, one of the extreme southern
stations for the species. Some of the San Jacinto Mountain specimens
120 University of California Publications in Zoology [ Vou. 17
are blotched and spotted like R. b. sierrae, and many show cross-bars
on the tibia. The skin is usually thickly studded with brownish, hispid
points, as in many boylii, but in the type it happens to be smooth. The
outer metatarsal tubercle is sometimes rudimentary. The head-width
enters the body-length from two and one-third to two and two-thirds
times.
Remarks.—The two very young specimens from Little Rock Creek
Canon show the light head patch of R. b. boylii, and the flexed tibia
extends beyond the anus as in that form. There is thus a good chance
that on the desert slope of the San Gabriel Mountains the frogs are
boylit rather than muscosa. The Phrynosoma blainvillii of the north-
ern San Gabriel Mountains (Pine Flats, Barley Flats, and the Upper
Tujunga Canon, 5000-5500 feet), and of the Sierra Liebre, is of the
northern subspecies, Ph. b. frontale; the frogs here discussed may offer
a parallel case.
Rana boylii muscosa inhabits the deeply cut valleys and gorges of
the San Gabriel, San Bernardino, and San Jacinto Mountains, from
at least the Arroyo Seco Canon near Pasadena, on the northwest, to
Keen’s Camp, Riverside County, on the southeast. It readily climbs
the steep rocks bordering the cafon streams, employing for this pur-
pose the enlarged tips of the digits, and sits far above the water during
the day; when alarmed it dives directly into the stream, kicks up the
silt with its hind legs, and buries itself in the mud, so that pursuit is
rendered difficult.
Rana boylii sierrae, new subspecies
Sierra Nevada Yellow-legged Frog
Type—Female, adult; no. 3734, Mus. Vert. Zool.; Matlack Lake,
10,500 feet altitude, two miles southeast of Kearsarge Pass, Sierra
Nevada, Inyo County, California; June 26, 1912; collected by H. 8S.
Swarth; orig. no. 9901.
Synonyms.—Rana aurora, part (Stejneger, 1893, p. 225); Rana
pretiosa, part (Stejneger, loc. cit., p. 226) ; [2] Rana pretiosa (Yarrow
and Henshaw, 1878, p. 1632), part [Lake Tahoe].
Diagnosis—With the general characters of Rana boyli boylii, but
hind leg usually shorter and head relatively narrower; tympanum
smoother ; and light patch on top of head wanting.
Description of type-—Vomerine teeth rudimentary, on two oblique
ridges nearly meeting between and slightly behind nares (see fig. 2) ;
1917] Camp: Notes on the Systematic Status of Toads and Frogs 121
head viewed from above rounded in outline; head-width contained
three times in body-length; hind limbs short, posterior side of bent
tarsus reaching forward to anterior corner of eye; fourth toe on hind
foot reaching forward to end of bent knee; dorso-lateral fold indis-
tinct, strongly pitted anteriorly ; tympanum nearly smooth, with seat-
tered hispid points; web of hind foot very large, extending to tips of
toes; outer metatarsal tubercle rudimentary, inner one small; plantar
tubereles small; tips of toes not much expanded; distal end of flexed
tibia, held at right angles to body, just reaching anus; color above, dark
yellowish brown, obseurely marked with indefinite darker vermicu-
lations ; lower surface yellow, faintly dotted with brown beneath chin ;
upper lip below eye mottled; no dark cheek patch; hind limbs not
distinetly barred with dark bands.
Material—One hundred and fifty specimens in the Museum of
Vertebrate Zoology from the following loealities in the Sierra Nevada
of California. Tulare County: Taylor Meadow, 7000 feet altitude
(nos. 8000-8010) ; Manter Meadow, 7000 feet (3012, 3013) ; Jackass
Meadow, 7750 feet (nos. 3014-3019) ; Monache Meadow (nos. 3020-
3033) ; vicinity of Ramshaw Meadow, 8800 feet (nos. 3034-3035) ;
Whitney Meadows, 9800 feet (nos. 3036-3053) ; Whitney Creek, 11,500
feet (no. 3055) ; Inyo County: Matlack Lake (as above) ; Yosemite
National Park: vicinity of Peregoy Meadow, 7000-7300 feet (nos.
781-5788, 5800) ; vicinity of Poreupine Flat, 8100 feet (nos. 5774,
775, 5789, 5803, 5804) ; Tenaya Lake (no. 5790) ; Tuolumne Meadows,
8600 feet (no. 5801) ; vicinity of Young Lake, 10,000 feet( nos. 5791-
5796) ; head of Lyell Canon, 9700—10,500 feet (nos. 5797-5799, 5802) ;
Vogelsang Pass, 10,450 feet (no. 6015) ; Vogelsang and Evelyn lakes,
10,850 feet (nos. 5962-5986, 5988-6014, 6016-6027); and Sunrise
Creek, 7300 feet (no. 5987).
Variations —Three specimens from the southern Sierra Nevada
and two from Young Lake, Tuolumne County, show faint traces of a
lighter patch across the anterior part of the head, as in R. b. boylii.
Nearly all the frogs from Whitney Meadows have the dorso-lateral
folds unusually well developed. The dorsal color pattern varies
widely. Occasionally the back is uniformly reddish-, brownish-, or
blackish-yellow; more rarely it is evenly marbled with dark brown
upon a dark yellow background with or without indefinite lighter
spots; more frequently the darker markings remain as indefinite
patches or as distinct black spots, thus approaching the pattern of
Rk. pretiosa. The outline of the head viewed from above is either
122 University of California Publications in Zoology [| Vou. 17
rounded as in the type or pointed as in boylii; in some specimens it 1s
as wide as in the narrowest boyli.
This seems to be the subspecies of boylii which approaches most
closely to the species pretiosa. A few specimens can be found in which
the vomerine teeth are confined to the ends of sight swellings on the
vomerine ridges, much as in pretiosa (see fig. 1); the outline of the
head is in some specimens the same as in pretiosa, and the short hind
leg carries the resemblance still farther. The dorsal color pattern,
while usually quite different from that in pretiosa, suggests the latter
in an occasional individual. I am inclined to place sierrae with boylw
on account of the character of the vomerine teeth in most of the speci-
mens (see figs. 1-3), the usual absence of distinct dorso-lateral folds,
the mottling of the upper lip, the lack of red in the coloration of the
Fig. 1—Inside of mouth of Rana pretiosa pretiosa, no. 5566, Mus. Vert. Zool.;
note vomerine teeth restricted to inner, expanded ends of oblique ridges, and
the large maxillary and premaxillary teeth.
Fig. 2.—Inside of mouth of Rana boylii sierrae, no. 3734, Mus. Vert. Zool.;
note resemblance of teeth to those of R. b. boylii.
Fig. 3—Inside of mouth of Rana boylii boylii, no. 6100, Mus. Vert. Zool.;
note vomerine teeth on whole length of oblique ridges and small teeth of upper
jaw. All X 114.
under parts, and the suggestion of a white, anterior head-patch in
seven or eight out of the 150 specimens examined.
Remarks.—This is the only species of frog known to occur above
7000 feet altitude in the Sierra Nevada. Its range is the entire south-
ern half of those mountains, at least from the Yosemite National Park
on the north to southern Tulare County on the south. It inhabits
meadows, streams and lakes from about 7000 to 10,500 feet in the
Yosemite Park, and to 11,500 feet near Mount Whitney. In some of
the lakes it was found in great numbers, appearing as soon as the ice
had melted in late June. The tadpoles were at this time of large size
and must have been hatched from eggs of a laying not more recent
than the previous year.
1917) Camp: Notes on the Systematic Status of Toads and Frogs 123
The specimen selected as the type is referred to by Mr. Harry 8.
Swarth in his field notes as having been the object of an attack by a
Clarke Crow. ‘‘The crow would peck at the frog and the latter would
squeal and puff himself out to twice his normal size. It was his cries
that attracted me.”’
KEY TO CALIFORNIA FROGS
1. Vomerine teeth rudimentary, on two oblique ridges between the nares (see
figs. 2, 3); tympanic region not darker than rest of head; fold along upper
lip colored like rest of body, mottled or dark; red never present in
coloration .. a Rana boylii and subspecies.
2. When hind leg is brought forward along body, inside angle of bent tarsus
and tibia reaching at least to nares and often beyond end of snout;
tympanum covered with many hispid points.
3. A light patch on top of head; darker area crossing the posterior half
of each upper eyelid merging insensibly into dorsal color behind;
body-length under 70 millimeters -................. Rana boylii boylii Baird.
3’. No light patch on top of head; darker areas crossing posterior half of
each upper eyelid, when present, contrasting with dorsal coloration;
hbody-length reaching 81 millimeters .......... Rana boylii muscosa Camp.
2’, When leg is brought forward, inside angle of bent tarsus seldom reaching
beyond nares; tympanum smooth or with but a few hispid points;
no light patch on top of head; body-length reaching 73 millimeters
Rana boylii sierrae Camp.
1’. Vomerine teeth large, in clusters on ends of indistinct oblique ridges inside
nares (see fig. 1); tympanic region darker than rest of head (except often
in pretiosa) ; fold along upper lip usually white or lighter than rest of
head; red often present in coloration.
4. When leg is brought forward along body, inside angle of bent tarsus
reaching to eye or nares, never beyond.
5. Back and top of head, set with inky, black spots; a small outer meta-
Gars allt CrCl es eecea eee cece Rana pretiosa pretiosa Baird and Girard.
5’. Baek, and top of head, with dark spots not so black as in pretiosa; no
outer metatarsal tubercle —........ Rana pretiosa luteiventris Thompson.
4’. Inside angle of bent tarsus and tibia, when brought forward, reaching to or
beyond nares; back and top of head without inky spots —..........-.-....
SSeS Pb E ces Bek ees Som see eee caer Fre ere Rae PP eer nee Tana aurora and subspecies.
6. Dorso-lateral folds indistinct; skin very smooth and thin; dorsal sur-
faces unspotted or with small dots; size medium; total length reaching
8 Ommaailiiimn(e ter Siescessce eee eee Rana aurora aurora Baird and Girard.
6’. Dorso-lateral folds prominent; skin thick and often slightly rough-
ened; dorsal surfaces with regularly placed, light centered spots;
size large; total length reaching 100 millimeters —.... nee Pee
Pes ee eee cee hse Rana aurora draytonii Baird and Girard.
124 University of California Publications in Zoology [ Vou. 17
Novres ON THE DISTRIBUTION OF WESTERN FROGS
Rana pretiosa pretiosa is known from nearly throughout Wash-
ington and Oregon, north into Canada, east into Montana, Wyoming,
and northern Utah, and south into extreme northern California.
Rana pretiosa luteiventris, a form recently described by Thompson
(1913, pp. 538-55) from Humboldt and Maggie valleys, Eureka and
Elko counties, Nevada, seems to be represented in the Museum of
Vertebrate Zoology by two specimens (nos. 2098, 2099), collected by
Dr. H. C. Bryant at Alturas, Modoe County, California.
Rana aurora aurora, described from Puget Sound, is known to the
present writer from the Willamette Valley, Oregon, and from Eureka,
Humboldt County, California, on the basis of specimens in the Museum
of Vertebrate Zoology. Specimens from Mendocino City, on the coast
of Mendocino County, California, are intergrades between aurora and
draytonn.
Rana aurora draytonii ranges north from the San Pedro Martir
Mountains, in Lower California, to Gualala, Mendocino County, Cali-
fornia (as indieated by material in the Museum of Vertebrate Zoology ),
and east from the coast to the foothills of the Sierra Nevada and into
the mountains of southern California up to an altitude of 4000 feet.
Transmitted October 28, 1916.
1917] Camp: Notes on the Systematic Status of Toads and Frogs 125
LITERATURE CITED
Batrp, 8. F.
1856. Descriptions of new genera and species of North American frogs.
Proe. Acad. Nat. Sci. Phila., 7, 1854-1855, 59-62.
Bairp, 8. F., and Girarp, C.
1854a. Descriptions of new species of reptiles, collected by the U. S. Ex-
ploring Expedition under the command of Capt. Charles Wilkes.
Proc. Acad. Nat. Sci. Phila., 6, 1852-1853, 174-177, 378-380.
1854b. List of reptiles collected in California by Dr. John L. LeConte, with
description of new species. Ibid., 300-302.
BOULENGER, G. A.
1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of
the British Museum. (Ed. 2; London), xvi + 503, 80 pls.
Corr, E. D.
1884. Notes on the geographical distribution of Batrachia and Reptilia in
western North America. Proe. Acad. Nat. Sci. Phila., 1883, 10-35.
1889. The Batrachia of North America. U.S. Nat. Mus. Bull., 34, 1-525,
86 pls.
DICKERSON, M. C.
1906. The frog book of North American toads and frogs, with a study of
the habits and life histories of those of the northeastern states.
(New York, Doubleday, Page & Co.), xvii + 253, 96 + 16 pls.
GIRARD, C.
1858. United States Exploring Expedition during the years 1838, 1839, 1840,
1841, 1842 under the command of Charles Wilkes, U.S. N. (Phila-
delphia, J. B. Lippincott & Co.), 20, Herpetology, xvii + 496; atlas,
pp- 1-10, 32 pls.
STEJNEGER, L.
1893. Annotated list of the reptiles and batrachians collected by the Death
Valley Expedition in 1891, with descriptions of new species. U.S.
Dept. Agric., Div. Ornithology and Mammalogy, N. Amer. Fauna,
7, 159-228, 394-398, 4 pls.
Tompson, H. B.
1913. Deseription of a new subspecies of Rana pretiosa from Nevada. Proce.
Biol. Soe. Wash., 26, 53-56, 1 pl.
Yarrow, H. C., and HensHaw, H. W.
1878. Report upon the reptiles and batrachians collected during the years of
1875, 1876, and 1877, in California, Arizona, and Nevada. Ann.
Rep. Chief U. 8. Engineers to Secy. of War., 1878, pt. 3 (Appen-
dices), Appendix L, in Appendix NN, pp. 1628-1648.
“zooLocy
Fok i
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UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 10, pp. 127-208, 14 figures in text July 11, 1917
A DISTRIBUTIONAL LIST OF THE AMPHIBIANS
AND REPTILES OF CALIFORNIA
BY
JOSEPH GRINNELL anp CHARLES LEWIS CAMP
(Contribution from the Museum of Vertebrate Zoology of the University of California)
The present paper deals with the systematic status and geographic
distribution of the amphibians and reptiles known to oeceur within the
present boundaries of the state of California. By including data
bearing on geographic distribution, and by placing the emphasis on
this rather than on the systematie phase of the subject, we hope to
forestall the criticism which is sometimes directed at a purely nominal
list. Restriction of the area treated to the limits of California has
made possible eritical study of most of the species included. We
believe that such a summary as is here presented is worth while at
the present time, in that it will accelerate inquiry in this field and
serve to direct the attention of workers towards the more conspicuous
gaps in our common knowledge.
The past two decades have witnessed the accumulation of much
information of the sort herewith presented. Many collections have
been made and a considerable amount of literature has appeared in
widely scattered places. Study of the large collection of newly gath-
ered specimens (6160) with full data now contained in the Museum
of Vertebrate Zoology and a critical review of the literature to date
form the basis of the present contribution.
The writers have followed Stejneger (Herpetology of Japan and
Adjacent Territory’) for the systematic arrangement used, and also
for most of the names of groups higher than genera. Where there
1U. S. Nat. Mus. Bull., 58, 1907, pp. xx + 577, 35 pls., 409 figs. in text.
128 University of California Publications in Zoology [Vou.17
has been question as to the applicability of specific names our decision
has been based on a precise interpretation of the rules of nomen-
clature and, where possible, upon recourse to type specimens. The
determination of species has been made at first hand, chiefly by the
junior author, on the basis of the collections and literature at hand,
with the exception of one genus: for the garter snakes (Thamnophis)
we have followed Ruthven (Variations and Genetic Relationships of
the Garter Snakes*).
The vernacular names chosen have been selected primarily upon
the grounds of previous use; sometimes fitness has been a factor in
their selection. The type locality of each species has been stated as
accurately as possible. Where the original statement of it was vague
we have attempted to supply a more exact designation. In some
cases this was not possible, and in a few others the type locality could
not be determined at all. Under ‘‘synonyms’’ and ‘‘common names’’
are included all names, both scientific and vernacular, other than the
accepted ones, which have been applied to the species as occurring
in California. Where the term ‘‘part’’ is used it means that the name
which it follows has been used for another Californian species. Where
a name now considered synonymous with an accepted name was based
on a specimen from California, the full citation and the type locality
are given; lack of space prevents giving the citations for all synonyms.
However, there is an index to all of these on file in the Museum of
Vertebrate Zoology, to which persons interested in following the sub-
ject farther are free to appeal. Needless to say, all of the specimens
in the Museum of Vertebrate Zoology are freely accessible for re-
examination by any student who may wish to verify our determi-
nations.
Besides the species and subspecies given regular place in the
present summary, all of which are known definitely to inhabit the
state, there are sixty or more species accredited to California on
grounds which do not seem to us satisfactory. Where possible we
have assigned these species to positions in the synonymy of forms in
good standing, but a number of them have had to be stricken entirely
from the list as indeterminable. We have not deemed it desirable to
list these numerous doubtful names here.
The statements of range for the several species involve the use of
three modes in which the distribution of vertebrate animals has been
designated. The first and more detailed statement is of a purely
2U. 8S. Nat. Mus. Bull., 61, 1908, pp. xii + 201, frontispiece pl., 82 figs. in text.
1917] Grinnell-Camp: Amphibians and Reptiles of California 129
geographic nature. Here, where the data permit, series of record
stations are given so as to outline roughly the range with regard to
political boundaries and towns, or to features of the topography and
drainage. Extreme points of occurrence on the north, east, south,
and west are thus given, sometimes several in each direction where a
long line is to be covered. For each of these the source of the record
is mentioned. This may be in published literature, in some one’s
unpublished notes, or from specimens in this museum.
Secondly, where practicable, the lfe-zone in which the species
occurs is given, for the purpose of bringing this paper into concordance
with others on the distribution of California vertebrates. Thirdly,
the habitat or associational restriction of the species is given. For
an explanation of these two latter modes of distributional designation
the reader is referred to a paper by the senior author (Grinnell, A
Distributional List of the Birds of California’).
The maps offered herewith serve to show the ranges of many of
the species of amphibians and reptiles in California from a purely
geographic standpoint. Stations of known occurrence are indicated
by appropriate signs, and with some of the species we have ventured
to indicate the outlines of the inferred ranges. By comparison of
these maps one with another and with maps showing the distribution
of birds and mammals in this state, a number of remarkable coinei-
dences in distribution will be observed. It must be that similar or,
possibly, the same factors operate to limit the distribution of such
widely different kinds of animals. Much yet needs to be realized in
the improvement of means of expressing the facts of animal distri-
bution; only a mere beginning has been made in systematizing such
knowledge as is already at hand. The meaning of distributional
limitation is one of the most fascinating subjects claiming the attention
of the field biologist. Really satisfactory inquiry into the underlying
causes must await the accumulation and assortment of vastly more
data than are here presented for reptiles and amphibians, and the same
degree of exhaustiveness must be applied to many different areas.
Most of the heretofore unpublished data used in the present paper
have been gathered during the past eight years for the Museum of
Vertebrate Zoology by various field parties working in its employ.
For the expense of bringing together and caring for the collections of
reptiles and amphibians now accessible in this museum, workers in
herpetology are indebted to Miss Annie M. Alexander. For the
3 Pacific Coast Avifauna, 11, 1915, 217 pp., 3 pls.
130 University of California Publications in Zoology (Vou. 17
privilege of examining California specimens in the United States
National Museum and for help in many questions of nomenclature the
present authors are indebted to Dr. Leonhard Stejneger, head curator
of biology in that institution. For help in the location of certain
geographic names and for critical assistance in proof-reading we owe
acknowledgment to Mr. Tracy I. Storer, of the staff of the Museum
of Vertebrate Zoology.
According to the present enumeration, there are represented in the
fauna of California one hundred and sixteen species and subspecies
of amphibians and reptiles. These fall into forty-six genera, twenty-
two families, and four orders.
Class AMPHIBIA
Order CAUDATA
Suborder MUTABILIA
Superfamily SALAMANDROIDEAE
Family SALAMANDRIDAE
Notophthalmus torosus (Rathke)
Pacifie Coast Newt
OriGiInaL Description.—Triton torosus Rathke, in Eschscholtz,
Zoologischer Atlas, pt. 5, 1833, pp. 12-14, pl. 21, fig. 15.
Type Locauiry.—Central California near coast, either at San
Francisco or between San Rafael, Marin County, and Fort Ross,
Sonoma County (see Kotzebue’s ‘‘New Voyage’’).
Synonyms.—Salamandra Beecheyi Gray, Zool. Beechey’s Voyage,
1839, p. 99, pl. 31, fig. 3 (type from Monterey, Monterey County) ;
“Triton Ermani Wiegmann, in Erman’s Reise um die Erde, Atlas,
1835, p. 24’’ [see Wiegmann’s Archive fiir Naturgeschichte, 2 Jahrg.,
2, 1836, pp. 163, 250] (type from California) ; Diemictylus torosus;
Taricha torosa; Molge torosa; Cynops torosus; Taricha laevis Baird
and Girard, Proe. Acad. Nat. Sci. Phila., 6, 1853, p. 302 (type from
San Francisco); Triton laevis; Amblystoma rubrum Reid, History
of Pasadena [California], 1895, p. 600 (type [by inference] from
near Pasadena) ; [?] Plewrodeles californiae; |?| Triton tereticauda
Eschscholtz, Zoologischer Atlas, pt. 5, 1833, p. 14 (type from Fort
Ross, Sonoma County) ; Aneides lugubris, part.
Common Names.—Western Newt; Warty Salamander ; Water-dog ;
Capt. Beechey’s Salamander; Pacific Water-lizard; California Newt;
Sad-colored Anaides, part.
1917] Grinnell-Camp: Amphibians and Reptiles of California 151
Rance.—Restricted to Pacifie drainage; most abundant west of
first coast divides. Found locally in Sacramento and San Joaquin
valleys, and in the central Sierra Nevada to 5500 feet altitude, as
near Cisco, Placer County (Mus. Vert. Zool.). Southernmost station,
a
Shes
Snes
iS 5
aS
Notophthalmus torosus
Ambystoma ensatum
Ambystoma tigrinum
Ambystoma paroticum
@oaopbppo
Ambystoma macrodactylum
Range of Notophthalmus torosus
outlined
2
Sr
DISTRIBUTION MAP
MUSEUM OF YERTEBRATE ZOOLOGY
Sy ~~
(eee NN =
Shares mes
UNIVERSITY OF CALIFORNIA
Fig. 1.
California.
Distribution of Salamanders (Notophthalmus and Ambystoma) in
San Diego River, San Diego County (Oreutt, West Amer. Scientist,
1, 1885, p. 5). Oceupies Upper Sonoran and Transition life-zones,
occasionally entering Canadian. Inhabits vicinity of streams and
ponds; in fall and winter the females live under rocks and logs, while
the males often remain in the water. (See fig. 1.)
132 University of California Publications in Zoology [Vou.17
Family PLETHODONTIDAE
Subfamily SPELERPINAE
Spelerpes platycephalus Camp
Mount Lyell Salamander
OrigiInaL Description.—Spelerpes platycephalus Camp, Univ.
Calif. Publ. Zool., 17, September 18, 1916, pp. 11-14, 5 figs.
Type Locaniry—Head of Lyell Cafion, 10,800 feet altitude, Yo-
semite National Park, Tuolumne County, California.
RanGceE.—Known only from the type locality, as above. This is
at the upper edge of the Hudsonian life-zone. The two specimens
were taken in a patch of heather among the rocks where water issued
from beneath surrounding snowbanks. (See fig. 2.)
Subfamily PLETHODONTINAE
Plethodon eschscholtzii (Gray)
Oregon Salamander
ORIGINAL DEscrIPTION.—Ensatina Eschscholtzii Gray, Cat. Am-
phibia Brit. Mus., pt. 2, 1850, p. 48.
Type Locaurry.—California: Monterey (fide Boulenger, Cat. Ba-
trachia Brit. Mus., ed. 2, 1882, p. 55).
Synonyms.—Plethodon ensatus; Plethodon oregonensis.
Common Names.—Oregon Triton; Oregon Plethodon.
RaNnGE.—Coast region. Has been taken south as far as Forest
Home, 5200 feet altitude, in the San Bernardino Mountains (no. 4890,
Mus. Vert. Zool.). Only one station interiorly from the coast belt:
[probably mountains near] Fresno (nos. 17,650-17,652, U. S. Nat.
Mus.). Oceurs in Transition and Upper Sonoran life-zones. Inhabits
damp, shady places in forests and heavy brush; often found beneath
masses of decaying vegetation and in holes in the earth. (See fig. 2.)
Plethodon croceater Cope
Yellow-spotted Salamander
OrigINAL Description.—Plethodon croceater Cope, Proc. Acad.
Nat. Sei. Phila., 1867 [1868], pp. 210-211.
Type Locauiry.—Fort Tejon, Kern County, California. (What
is probably the type is contained in the collection of the Department
of Zoology, University of California.)
1917] Grinnell-Camp: Amphibians and Reptiles of California 183
Synonym.—|[?] Plethodon flavipunctatus Strauch, Mem. Acad.
Imper. Sci. St. Petersb., ser. 7, 16, no. 4, 1870, p. 71 (type from ‘*‘ New
Albion,’’ California) [see Van Denburgh, Proe. Calif. Acad. Scei.,
ser. 4, 6, 1916, p. 221].
Aneides lugubris lugubris
Aneides lugubris farallonensis
Aneides ferreus
Aneides iécanus
Plethodon eschscholtzti
Plethodon croceater
Plethodon elongatus |
Spelerpes platycephalus
+rPLC TEBE a0 e@eO
Ranges of Aneides lugubris lugubris
and Plethodon croceater outlined
N
iN
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA
—
Fig. 2. Distribution of Salamanders (Plethodon, Spelerpes, and Aneides) in
California.
Common Names.—Cape St. Lucas Triton; Yellow-spotted Lizard.
RanGE.—Sierra Nevada Mountains (Van Denburgh, loc. cit.).
Definite stations of occurrence are: Mud Spring, 6300 feet altitude,
4 miles southwest of Nelson, Tulare County (3 specimens, Mus. Vert.
Zool.) ; Grant Forest, Sequoia National Park, Tulare County (2 speci-
134 University of California Publications in Zoology [Vou.17
mens, Univ. Calif. Dept. Zool.) ; Alta, 3600 feet altitude, Placer County
(1 specimen, Univ. Calif. Dept. Zool.); Fort Tejon (as above).
Occurs within the Transition life-zone. Inhabits damp places in
forests. (See fig. 2.)
Plethodon elongatus Van Denburgh
Del Norte Salamander
OriewaL DEscription.—Plethodon elongatus Van Denburgh, Proe.
Calif. Acad. Sci., ser. 4, 6, May 12, 1916, pp. 216-218.
Type Locaniry.—Requa, Del Norte County, California.
Rance.—Known only from the type locality, as above. (See fig. 2.)
Aneides lugubris lugubris (Hallowell)
Arboreal Salamander
Original DescripTion.—Salamandra lugubris Hallowell, Proc.
Acad. Nat. Sei. Phila., 4, 1848 [1850], p. 126.
Type Locauiry.—Monterey, California.
Synonyms.—Anaides lugubris, part; Ambystoma punctulatum
Gray, Cat. Batr. Caud. in Brit. Mus., 1850, p. 37 (type from Monte-
rey) ; Ambystoma punctatum, part; Taricha lugubris; Autodax lugu-
bris, part; Plethodon crassulus Cope, Proce. Amer. Philos. Soe., 23,
1886, pp. 521-522 (type from California) [Van Denburgh (Proc.
Calif. Acad. Sci., ser. 4, 6, 1916, pp. 219-220) regards this name as a
synonym of Plethodon intermedius, and doubts that the type really
came from California].
Common Names.—California Land Salamander; Sad-colored
Anaides, part; Spotted Autodax; Mournful Salamander; Speckled
Salamander.
Rance.—Abundant in the inner coast region. Occurs north as
far as Mendocino City, Mendocino County (Mus. Vert. Zool.). The
two isolated interior stations are: 3 miles northeast of Coulterville,
3100 feet altitude, Mariposa County, and Los Banos, Mereed County
(Mus. Vert. Zool.). Southeasternmost occurrence within the state:
San Diego County (U.S. Nat. Mus.; also see Miller, Amer. Nat., 40,
1906, pp. 741-742). Oceurs chiefly in the Upper Sonoran life-zone,
though extending into adjacent Transition and into Lower Sonoran.
Lives in hollows of oak trees and also on the ground beneath stones
and logs, as a rule in oak woods. (See fig. 2.)
1917] Grinnell-Camp: Amphibians and Reptiles of California 135
Aneides lugubris farallonensis (Van Denburgh)
Farallon Salamander
OrigtnaL Description.—Autodaxr lugubris farallonensis Van Den-
burgh, Proe. Calif. Acad. Sei., ser. 3, zool., 4, June 15, 1905, pp. 5-6,
pl. 2.
Type Locauiry.—South Farallon Island, San Francisco County,
California.
Synonyms.—A naides lugubris, part; Autodax lugubris, part.
Rance.—Known only from South Farallon Island. Found under
rocks. (See fig. 2.)
Aneides ferreus (Cope)
Rusty Salamander
OrietnaL DeEscripTion.—Anaides ferreus Cope, Proce. Acad. Nat.
Sei. Phila., 1869, pp. 109-110.
Type Locaniry.—Fort Umpqua, Douglas County, Oregon.
RaNGE.—Extreme northern humid coast belt. Recorded only from
the following localities (Van Denburgh, Proe. Calif. Acad. Sci., ser. 4,
6, 1916, p. 216): Requa, Del Norte County; Alton, Trimidad, and
Carlotta, Humboldt County; Comptche, Mendocino County. These
localities le in the Transition life-zone. Specimens at Requa were
taken from the rotten wood of a dead tree, twenty feet above the
ground. (See fig. 2.)
Aneides iécanus (Cope)
Shasta Salamander
ORIGINAL DESCRIPTION.
Sci. Phila., 1883, pp. 24-25.
Type Locauiry.—Baird, Shasta County, California.
Plethodon vécanus Cope, Proce. Acad. Nat.
Synonyms.—A naides vécanus; Autodaxr vécanus.
Common Name.—Black Salamander.
Raner.—Known only from west-central and northern California,
the following being the record stations known to date: Baird and
McCloud River, Shasta County (Townsend, Proce. U. S. Nat. Mus., 10,
1887, pp. 240-241) ; Humboldt County (Univ. Calf. Dept. Zool.) ;
Ukiah, Mendocino County (Mus. Vert. Zool.) ; Los Gatos and Stevens
Creek, Santa Clara County, and Glenwood and Boulder, Santa Cruz
136 University of California Publications in Zoology [Vou.17
County (Van Denburgh, Proc. Calif. Acad. Sci., ser. 2, 5, 1895, pp.
776-778) ; Mountain View, Santa Clara County (no. 21170, U.S. Nat.
Mus.). Oceurs mainly within the Transition life-zone. Lives in
recesses In the ground and beneath logs, stones, and boards, often in
the vicinity of running water. (See fig. 2.)
Batrachoseps pacificus (Cope)
Island Salamander
OrieinaAL DescripTion.—Hemidactylium pacificum Cope, Proce.
Acad. Nat. Sei. Phila., 1865, pp. 195-196.
Type Locauiry.—Santa Barbara, California [doubtless one of the
Channel Islands near Santa Barbara: Van Denburgh, Proc. Calif.
Acad. Sci., ser. 3, zool., 4, 1905, p. 7].
Common Name.—Pacifie Lizard.
Rance.—Known only from San Miguel, Santa Rosa, and Santa
Cruz islands, off the coast of southern California (Van Denburgh,
Proe. Calif. Acad. Sci., ser. 4, 4, 1914, pp. 1384, 135). (See fig. 3.)
Batrachoseps major Camp
Garden Salamander
ORIGINAL Descrietion.—Batrachoseps major Camp, Uniy. Calif.
Publ. Zool., 12, April 2, 1915, pp. 827-330.
Type Locaurry.—Sierra Madre (the town), 1000 feet altitude,
Los Angeles County, California.
SynonyM.—[?] Batrachoseps attenuatus, part.
Rance.—Known only from the Pacifie slope of southern California,
where recorded definitely only from Pasadena, and from the type
locality, as above. Occurs within the Lower Sonoran life-zone. In
the rainy season lives at the surface of the ground beneath boards and
stones; in the dry season retreats into the earth. (See fig. 3.)
Batrachoseps attenuatus (Eschscholtz)
Slender Salamander
ORIGINAL DescriptTiIoN.—Salamandrina attenwata Eschscholtz, Zo-
ologischer Atlas, pt. 5, 1833, pp. 1-6, pl. 21.
Type Locaniry.—Central California near coast, either at San
Francisco or between San Rafael, Marin County, and Fort Ross,
Sonoma County.
1917] Grinnell-Camp: Amphibians and Reptiles of California 137
Synonyms.—Salamandra attenuata; |?) Batrachoseps pacificus,
part; Batrachoseps nigriventris Cope, Proce. Acad. Nat. Sei. Phila.,
1869, p. 98 (type from Fort Tejon, Kern County).
Common Names.—Slender Lizard; Blaek-bellied Lizard.
o Batrachoseps attenuatus
@ Batrachoseps major
ov Batrachoseps pacificus
Range of Batrachoseps attenuatus
outlined
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALLPORNLA
Ranee.—Coast district from Ukiah, Mendocino County, south at
least to mountains immediately north of Claremont, Los Angeles
County; occurs also along the foothills of the central Sierra Nevada,
north to Eldorado County (U.S. Nat. Mus.) and southeast to three
miles south of Nelson, Tulare County (Mus. Vert. Zool.), and on Santa
Catalina Island (Van Denburgh, Proce. Calif. Acad. Sci., ser. 3, zool.,
138 University of California Publications in Zoology (Vou. 17
4, 1905, p. 16). Oceurs chiefly within the Upper Sonoran life-zone,
though entering Transition and upper edge of Lower Sonoran }oeally.
Found in rotten logs, in damp masses of dead vegetation, beneath
rocks on shaded slopes, and in rodent burrows in the ground. (See
fig. 3.)
Family AMBYSTOMIDAE
Ambystoma tigrinum (Green)
Tiger Salamander
OrieInAL Description.—Salamandra tigrina Green, Journ. Acad.
Nat. Sei. Phila., 5, 1825, pp. 116-118, pl. 25, fig. 7.
Typr Locaniry—Near Moorestown, New Jersey.
Synonyms.—Ambystoma tigrinum, part; Ambyostoma califor-
niense Gray, Proe. Zool. Soe. Lond., 21, 1853, p. 11, pl. 7 (type from
Monterey) ; Ambystoma tigrinuwm califormense; Ambystoma mavor-
tium californiense; Ambystoma mavortium; [?] Ambystoma ingens;
[?] Heterotriton ingens; Amblystoma californicum.
Common Names.—California Axolotl; California Salamander.
Rancu.—tThe west-central part of the state, where known from the
following localities: Galt, Sacramento County; Ripon, San Joaquin
County; and Mount Hamilton, Santa Clara County (Storer, Copeia,
no. 24, November, 1915, p. 56); Monterey, Monterey County (as
above); Fresno, Fresno County; Fort Tejon. Kern County; and
Petaluma, Sonoma County (Cope, U. 8S. Nat. Mus. Bull., 34, 1889,
pp. 85-86). These loealities lie in the Upper Sonoran life-zone, save
for Fresno, which is in the Lower Sonoran, and Monterey, which is
in Transition. Inhabits burrows of rodents in open country. (See
fig. 1.)
Ambystoma macrodactylum Baird
Long-toed Salamander
OrIGINAL DeEscripTiION—Ambystoma macrodactyla Baird, Journ.
Acad. Nat. Sei. Phila., ser. 2, 1, 1850, p. 292.
Type Locaniry.—Astoria, Oregon.
Common Name.—F lat-footed Salamander.
Rance.—High northeastern section of the state, with record
stations as follows: ‘‘California’’ (Boulenger, Cat. Batr. Brit. Mus.,
ed. 2, 1882, p. 48) ; near Medicine Lake, 6500 feet altitude, Siskiyou
1917] Grinnell-Camp: Amphibians and Reptiles of California 139
County (Univ. Calif. Dept. Zool.) ; near Fallen Leaf Lake, Eldorado
County (Van Denburgh, Proe. Calif. Acad. Sci., ser. 4, 6, 1916, p. 215).
Life-zone, Transition. (See fig. 1.)
Ambystoma ensatum (Eschscholtz)
Marbled Salamander
OriagInaAL Description.—Triton ensatus Eschscholtz, Zoologischer
Atlas, pt. 5, 1833, pp. 6-12, pl. 22.
Type Locaurry.—Central California near coast, possibly near Fort
Ross, Sonoma County.
Synonyms.—Chondrotus tenebrosus; Amblystoma tenebrosum;
Dicamptodon ensatus; Xiphonura tenebrosa; Chondrotus lugubris.
Common NameE.—Oregon Salamander, part.
RaneGe.—Chiefly the humid coast belt, occurring south as far as
six miles north of Boulder, Santa Cruz County (Camp, MS). Re-
corded also from MeCloud River, Shasta County (Townsend, Proce.
U. S. Nat. Mus., 10, 1887, p. 240), and Allen Springs, Lake County
(Univ. Calif. Dept. Zool.). Oceupies the Transition life-zone. In-
habits tracts of forest and heavy brush; lives in streams, or under
rocks and in holes in wet earth. (See fig. 1.)
Ambystoma paroticum Baird
British Columbia Salamander
Oriana Description.—A mblystoma paroticum Baird, Proe. Acad.
Nat. Sei. Phila., 1867 [1868], pp. 200-201.
Type Locaurry.—Chiloweyuek, Washington Territory [== Chilli-
wack Lake, British Columbia].
Common Nami.—Vancouver’s Salamander.
Rance—Recorded only from Requa, Del Norte County (Van
Denburgh, Proc. Calif. Acad. Sci., ser. 4, 6, 1916, p. 216). Found in
wet earth under a stump. (See fig. 1.)
Order SALIENTIA
Suborder COSTATA
Family DISCOGLOSSIDAE
Ascaphus truei Stejneger
American Bell-toad
ORIGINAL DescripTion.—Ascaphus truei Stejneger, Proe. U. 8.
Nat. Mus., 21, June 20, 1899, pp. 899-901, pl. 89.
140 University of California Publications in Zoology (Vou. 17
Type Locaniry.—Humptulips, Chehalis County, Washington.
RanGEe.—One capture: Northwest slope Craggy Peak, elevation
5000 feet, Siskiyou Mountains, Siskiyou County (no. 45362, U.S. Nat.
Mus.).
Suborder LINGUATA
Family PELOBATIDAE
Scaphiopus hammondii hammondii Baird
Western Spadefoot
OrIGINAL DEscriIpTION.—Scaphiopus hammondii Baird, Pac. R. R.
Rep., 10, 1859, Abbot’s report, pt. 4, no. 4, p. 12, pl. 28 (figs. 2a, 2b,
2c, 2d).
Type Locauiry.—Fort Reading [near Redding, in Shasta County],
California.
Synonyms.—Spea hammondu; Spea stagnalis.
Common Names.—Hammond’s Spea; New Mexican Spea; Ham-
mond’s Spadefoot.
RaNncE.—Semi-arid portions of the state both east and west of the
Sierra Nevada, from San Diego (Cooper, in Cronise, Nat. Wealth
Calif., 1868, p. 486) to Fort Reading, as above. Present records
include: Los Angeles (Univ. Calif. Dept. Zool.) and Sierra Madre
(Mus. Vert. Zool.), Los Angeles County ; Santa Barbara (Yarrow and
Henshaw, Ann. Rep. U. 8. Engineers, 1878, p. 1631) ; Olancha, Inyo
County (Stejneger, N. Amer. Fauna, 7, 1893, p. 222); Benton and
Mono Lake, Mono County (Mus. Vert. Zool.) ; Lane’s Bridge, Fresno
County (Mus. Vert. Zool.) ; and Simmler, San Luis Obispo County
(Mus. Vert. Zool.). Occurs chiefly in the Lower Sonoran life-zone,
but extends also into Upper Sonoran. Inhabits open country where
soil is of a nature to permit burrowing to safe depths during dry
periods.
Family BUFONIDAE
Bufo cognatus cognatus Say
Great Plains Toad
OrIGINAL Description.—Bufo cognatus Say, in Long’s Expedition
to Rocky Mountains, 2, 1823, p. 190, footnote.
Type Locatiry.—Arkansas River [in Colorado, probably between
present site of La Junta and the Colorado-Kansas boundary].
1917] Grinnell-Camp: Amphibians and Reptiles of California 141
Rance.—Only along the Colorado River north to Needles, San
Bernardino County, and in the Salton Basin northwest to Mecea,
Riverside County (Mus. Vert. Zool.). Life-zone, Lower Sonoran.
Inhabits overflow bottom-lands and irrigated districts. (See fig. 4.)
Bufo cognatus cognatus
Bufo cognatus californicus
Bufo boreas boreas
Bufo boreas halophilus
Intergrade
Bufo boreas nelsoni 4
Bufo canorus
Bufo punctatus
Bufo alvarius |
Bufo woodhousii
+pP1LB880000e
Range of Bufo boreas and
subspecies solidly outlined
Southern limits of Bufo boreas
boreas indicated by broken line
DISTRIBUTION MAP
MUSEUM OF YERTEBRATE ZOOLOGY
Fig. 4. Distribution of Toads (Bufo) in California.
Bufo cognatus californicus Camp
Arroyo Toad
Ortemnau Description.—Bufo cognatus californicus Camp, Univ.
Calif. Publ. Zool., 12, April 2, 1915, pp. 331-334.
Type Locauiry.—Santa Paula, Ventura County, California.
142 University of California Publications in Zoology [Vou.17
RanceE.—Known only from the type locality, as above, and from
the Tujunga Wash, near Sunland, Los Angeles County, California
(Camp, loc. cit.). Both these places he in the Lower Sonoran life-
zone. Found in vicinity of dry washes. (See fig. 4.)
Bufo woodhousii Girard
Roeky Mountain Toad
ORIGINAL DEscrIPTION.—Bufo woodhousti Girard, Proe. Acad. Nat.
Sci. Phila., 7, 1854, p. 86 (this name to replace Bufo dorsalis of Hallo-
well, Proce. Acad. Nat. Sei. Phila., 6, 1852, pp. 181-182).
Type Locauiry.—New Mexico [= Arizona; type no. 2531 in U.S.
Nat. Mus., labeled ‘‘Calif. Mountains, Mexico,’’ which equals San
Francisco Mountains, Arizona: see Stejneger, N. Amer. Fauna, 3, 1890,
pp. 116-117].
Synonyms.—Bufo lentiginosus americanus; Bufo lentiginosus
woodhousu.
Common Names.—Woodhouse’s Toad; American Toad.
Rance.—Along the Colorado River (Cooper, in Cronise, Nat.
Wealth Calif., 1868, p. 486). Stations of recent capture are: near
Potholes and 5 miles northeast of Yuma, Imperial County (Mus. Vert.
Zool.). Both stations are on the bottom-lands and in the Lower
Sonoran life-zone. (See fig. 4.)
Bufo boreas halophilus Baird and Girard
California Toad
OriGiInaL Descrrprion.—Bufo halophila Baird and Girard, Proe.
Acad. Nat. Sei. Phila., 6, 1853, p. 301.
Tyre Locauiry.—Benicia, Solano County, California.
Synonyms.—Bufo boreas, part; Bufo chilensis; Bufo columbiensis,
part; Bufo columbiensis halophilus; Bufo borcas nelsoni, part.
Common Names.—Baird’s Toad, part; Common Toad; Salt-marsh
Frog.
Rance—Throughout both the coast region and the interior valleys,
north typically to Gualala River, Sonoma County, to Oroville, Butte
County, and to vicinity of Independence, Inyo County ; east, centrally,
into Owens Valley, and to Antelope Valley, Los Angeles County, and
Victorville, San Bernardino County; east at the south to Vallecito
and La Puerta, San Diego County (Mus. Vert. Zool.). Southernmost
1917] Grinnell-Camp: Amphibians and Reptiles of California 143
station, San Diego (Yarrow, U. 8. Nat. Mus. Bull., 24, 1882, pp. 23,
162). Absent from deserts bordering Colorado River. Occurs in all
life-zones below upper edge of Hudsonian. Inhabits open valleys and,
rarely, wooded areas; in the high mountains inhabits wet meadows and
lake shores; lives during the day in gopher and ground-squirrel bur-
rows and under rocks. (See fig. 4.)
Bufo boreas boreas Baird and Girard
Northwestern Toad
OriegInAL DescripTion.—Bufo boreas Baird and Girard, Proce.
Acad. Nat. Sei. Phila., 6, 1852, pp. 174-175.
Type Locauiry.—Columbia River and Puget Sound.
Synonyms.—Bufo columbiensis, part; Bufo halophilus, part; Bufo
microscaphus.
Common Names.—Baird’s Toad, part; Small-spaded Toad.
Rance.—Northern portion of the state, south to Eureka, Humboldt
County, to Sisson, Siskiyou County, and to Mono County (Mus. Vert.
Zool.). From these localities and others adjacent, specimens show
intergradation towards B. b. halophilus. Occurs in the Transition
and Upper Sonoran life-zones. Lives in tules about lake shores, along
streams, and in mountain meadows. (See fig. 4.)
Bufo boreas nelsoni Stejneger
Nevada Toad
OrienaAL Description.—Bufo boreas nelsoni Stejneger, N. Amer.
Fauna, 7, May 31, 1893, pp. 220-221, pl. 3, figs. 4a, 4b.
Type Locauiry.—Oasis Valley, Nye County, Nevada.
Rance.—Eastern Inyo region. Recorded from Resting Springs,
Inyo County (Stejneger, loc. cit.). Life-zone, Lower Sonoran. (See
fig. 4.)
Bufo canorus Camp
Yosemite Park Toad
OrigInaL DeEscripTion.—Bufo canorus Camp, Univ. Calif. Publ.
Zool., 17, November 17, 1916, pp. 59-62, figs. 1-4.
Type Locauiry.—Porcupine Flat, 8100 feet altitude, Yosemite
National Park, Mariposa County, California.
RanGe.—High central Sierra Nevada, 7000 to 11,000 feet altitude,
almost altogether within the limits of the Yosemite National Park
144 University of California Publications in Zoology [|Vou.17
(Mus. Vert. Zool.). Oceupies the Canadian and Hudsonian life-zones,
extending even into Alpine-Arctic. Inhabits vicinity of meadows and
the margins of streams and lakes. (See fig. 4.)
Bufo punctatus Baird and Girard
Spotted Toad
OriegInAL DEscriPTION.—Bufo punctatus Baird and Girard, Proe.
Acad. Nat. Sci. Phila., 6, 1852, p. 173.
Type Locauiry.—Rio San Pedro of the Rio Grande del Norte
(in Texas?).
Synonym.—Bufo beldingi.
Rance.—Colorado desert, west to Vallecito, San Diego County, and
Carrizo Creek, north slope Santa Rosa Mountains, Riverside County
(Mus. Vert. Zool.) ; also Turtle Mountains, near Blythe Junction, San
Bernardino County (Camp, Univ. Calif. Publ. Zool., 12, 1916, p. 512) ;
Furnace Creek, Death Valley, and Cottonwood Canon [Jackass
Spring], Panamint Mountains, in Inyo County (Stejneger, N. Amer.
Fauna, 7, 1893, p. 219). Life-zone, Lower Sonoran. Inhabits rocky
canon bottoms, near springs or streams. (See fig. 4.)
Bufo alvarius Girard
Colorado River Toad
OriGINaAL DerscripTion.—Bufo alvarius Girard, in Baird, U. S.
Mex. Bound. Surv., 2, 1859, pt. 2, reptiles, p. 26, pl. 41, figs. 1-6.
Type Locauiry—Valley of Gila and Colorado: Fort Yuma, Cali-
fornia (see Cope, U. S. Nat. Mus. Bull., 34, 1889, p. 267).
Common Names.—Girard’s Toad; Colorado Toad.
Rance.—Immediate valley of the Colorado River. Has been taken
in the delta region west to Meloland (Mus. Vert. Zool.) ; all known
localities for the state are in Imperial County. Life-zone, Lower
Sonoran. Inhabits only riparian bottom-lands and irrigated districts.
(See fig. 4.)
Family HYLIDAE
Subfamily Hyninar
Hyla regilla Baird and Girard
Pacific Tree-frog
OriGgInAL Description.—Hyla regilla Baird and Girard, Proe.
Acad. Nat. Sei. Phila., 6, 1852, p. 174.
1917] Grinnell-Camp: Amphibians and Reptiles of California 145
Type Locauiry.—Sacramento River, California.
Synonyms.—Litoria occidentalis Baird and Girard, Proe. Acad.
Nat. Sei. Phila., 6, 1853, p. 301 (type from San Francisco) ; Hyla
scapularis; Hyla nebulosa Hallowell, Proce. Acad. Nat. Sei. Phila., 7,
1854, pp. 96-97 (type from Tejon Pass, probably near Fort Tejon,
Kern County) ; Hyla scapularis var. hypochondriaca Hallowell, Proce.
Acad. Nat. Sci. Phila., 7, 1854, p. 97 (type from Tejon Pass) ;
Hyla cadaverina (a name to replace nebulosa of Hallowell: Cope,
Journ. Acad. Nat. Sci. Phila., ser. 2, 6, 1866, p. 84) ; Hyla regilla var.
scapularis.
Common Names.
Western Tree-frog; Wood-frog; Pacifie Hyla;
Tree-toad, part; Cadaverous Hyla; Greeny; Cape San Lueas Hyla.
Rance.—AImost throughout the state, except in the extreme south-
eastern desert district. Occurs on Santa Rosa, Santa Cruz, and Santa
Catalina islands (Van Denburgh, Proe. Calif. Acad. Sci., ser. 4, 4,
1914, pp. 182, 135, 137). Extends to timber-line in the Sierra Nevada;
occurs in all zones below Alpine-Arctic. Inhabits damp recesses among
rocks and logs; the ground in the vicinity of springs, streams, and
lakes; rank growths of vegetation, especially in marshy places; trees
in damp forests; and, in open country, burrows of various animals.
Hyla arenicolor Cope
Arizona Tree-frog
Originau Description.—Hyla arenicolor Cope, Journ. Acad. Nat.
Sci. Phila., ser. 2, 6, 1866, p. 84 (a name to replace Hyla affinis, Baird,
Proce. Acad. Nat. Sei. Phila., 7, 1854, p. 61).
Type Locauiry.—Northern Sonora [Mexico or Arizona?].
Synonyms.—Hyla copii; Hyla versicolor.
Common Names.—Cope’s Tree-frog; Tree-toad, part ; Cope’s Hyla;
Sand-colored Tree Frog.
Rance.—Northwest from Julian, La Puerta, and Mountain Spring,
in eastern San Diego County, to near Nordhoff, Ventura County, and
Santa Monica Mountains, Los Angeles County; east to Little Rock
Creek, Los Angeles County, and to Banning, Dos Palmos Spring, and
Deep Canon, Riverside County; also on Pine Mountain, near Escon-
dido, San Diego County (Mus. Vert. Zool.; Richardson, Amer. Nat.,
46, 1912, pp. 605-611). Oceupies chiefly the Upper Sonoran life-zone,
extending locally into Lower Sonoran. Lives on boulders and exposed
rock faces close to cafion streams.
146 University of California Publications in Zoology [Vou.17
Family RANIDAE
Subfamily RanmagE
Rana boylii boylii Baird
California Yellow-legged Frog
OricInAL Description.—Rana boylii Baird, Proce. Acad. Nat. Sci.
Phila., 7, 1854, p. 62.
Tyre Locauiry.—California; subsequently designated as El Do-
rado [Eldorado County ?] (see Cope, U. S. Nat. Mus. Bull., 34, 1889,
p. 447).
Synonyms.—Rana pachyderma Cope, Proce. Acad. Nat. Sei. Phila.,
1883, pp. 25-27 (types from MeCloud River [= Baird, Shasta
County], California, and Ashland, Oregon); [?] Rana temporaria
pretiosa, part.
Common Names.—Thick-skinned Frog; Boyle’s Frog.
RanGe.—Northern and central portions of the state, chiefly west
of the high Sierra Nevada, both along the coast and in the interior
valleys. Has been found east to Baird, Shasta County (Cope, loc. cit.),
to Fyffe, Eldorado County (Mus. Vert. Zool.), and to near Feliciana
Mountain, 3800 feet altitude, Mariposa County (Mus. Vert. Zool.).
Southernmost station, Bodfish, on Kern River, Kern County (Mus.
Vert. Zool.). Has also been taken at Mono Lake, Mono County (Mus.
Vert. Zool.). Life-zones, Upper Sonoran and Transition. Inhabits
margins of springs, streams, and fresh-water lakes. (See fig. 5.)
Rana boylii sierrae Camp
Sierra Nevada Yellow-legged Frog
ORIGINAL DescripTion.—Rana boylii sierrae Camp, Univ. Calif.
Publ. Zool., 17, February 3, 1917, pp. 120-123, fig. 2.
Type Locauiry.—Matlack Lake, 10,500 feet altitude, near Kear-
sarge Pass, Sierra Nevada, Inyo County, California.
Synonyms.—Rana aurora, part; Rana pretiosa, part; [?] Rana
temporaria pretiosa, part.
Common Names.—Western Frog; Pacific Frog.
Rance.—High Sierra Nevada, 7000 to 10,500 feet altitude. North-
ernmost station, Young Lake, Yosemite National Park, Tuolumne
County ; southernmost, Taylor Meadow, near Kern County line, Tulare
County (Mus. Vert. Zool.). Oceupies the Canadian and Hudsonian life-
zones. Lives chiefly in lakes and along streams in meadows. (See fig. 5.)
1917] Grinnell-Camp: Amphibians and Reptiles of California 147
Rana boylii muscosa Camp
Sierra Madre Yellow-legged Frog
OrieinaL Descrrerion.—Rana boylii muscosa Camp, Univ. Calif.
Publ. Zool., 17, February 3, 1917, pp. 118-120.
Rana boylit boylii
Rana boylii muscosa
Rana boylit sierrae
Rana pretiosa pretiosa
Rana pretiosa luteiventris
Rana aurora aurora 4
Rana aurora draytonii
Intergrade
+16) (O10) (Pp on) Een
Rana pipiens
Ranges of Rana boylii and
Rana aurora outlined
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OP CAJ.IFORNLA Ss
|
Fig. 5. Distribution of Frogs (Rana) in Califernia.
Type Locauiry—Arroyo Seco Canon, near Pasadena, California.
Rance—San Gabriel (both the desert and Pacific drainages), San
Bernardino and San Jacinto mountains. Northwesternmost stations
for typical specimens, Arroyo Seco Cafion near Pasadena, and Little
Rock Creek, in Los Angeles County ; southeasternmost station, Straw-
148 University of California Publications in Zoology [Vou.17
berry Valley, Riverside County (Mus. Vert. Zool.). Oceupies the
Upper Sonoran and Transition life-zones. Lives along streams in
narrow rock-walled canons. (See fig. 5.)
Rana pretiosa pretiosa Baird and Girard
Western Spotted Frog
OrIGINAL DEscripTiIoN.—Rana pretiosa Baird and Girard, Proe.
Acad. Nat. Sci. Phila., 6, 1853, p. 378.
Type Locauiry.—Puget Sound.
Rance.—Extreme northern end of the state. Definite stations of
occurrence are Fall City Mills, Fall River, Shasta County (U.S. Nat.
Mus., no. 38806), and vicinity of Sisson, Siskiyou County (Mus. Vert.
Zool. and U.S. Nat. Mus.). Life-zone, Transition. Inhabits marshes
and lakes. (See fig. 5.)
Rana pretiosa luteiventris Thompson
Nevada Spotted Frog
OrIGINAL DeEscription.—Rana pretiosa luteiventris Thompson,
Proe. Biol. Soe. Wash., 26, March 22, 1913, pp. 53-55, pl. 3, figs. 2, 3.
Type Locaniry.—Anne Creek, Elko County, Nevada.
Rance.—Eastern Modoe County. Only station, Pine Creek, near
Alturas, Modoe County (Mus. Vert. Zool., nos. 2098, 2099). Speci-
mens captured on banks of a slow-flowing stream. (See fig. 5.)
Rana aurora aurora Baird and Girard
Oregon Red-legged Frog
ORIGINAL DeEscription—Rana aurora Baird and Girard, Proce.
Acad. Nat. Sei. Phila., 6, 1852, p. 174.
Type Locaniry.—Puget Sound.
Rance.—Extreme northwestern corner of the state. Stations of
capture all in vicinity of Eureka, Humboldt County (Mus. Vert.
Zool.). (See fig. 5.)
Rana aurora draytonii Baird and Girard
California Red-legged Frog
OriGgInAL Description.—Rana Drayton Baird and Girard, Proe.
Acad. Nat. Sei. Phila., 6, 1852, p. 174.
Type Locaniry.—San Francisco, California.
Synonyms.—Rana Lecontii Baird and Girard, Proe. Acad. Nat.
Sci. Phila., 6, 1853, pp. 301-302 (type from San Francisco) ; Rana
1917] Grinnell-Camp: Amphibians and Reptiles of California 149
nigricans Hallowell, Proe. Acad. Nat. Sci. Phila., 7, 1854, p. 96 (type
from El Paso Creek, Kern County, California) ; Rana longipes Hallo-
well, Pac. R. R. Rep., 10, 1859, Wilhamsons’ Rep., pt. 4, no. 1, pp.
20-21 (type from El Paso Creek, Kern County, California) ; Epirhexis
longipes; Rana agilis aurora; Rana catesbiana; Rana septentrionalis ;
Rana temporaria; Rana temporaria aurora.
Common Names.—Drayton’s Frog; Bull-frog; Long-footed Frog ;
Bloody Nouns; Rocky Mountain Frog; Western Wood Frog; Leconte’s
Frog.
Ranee.—Central and southern portions of the state, west of the
Sierran divides. Extends from the seaboard east to about 3500 feet
altitude on the west slope of the central Sierra Nevada. Northern
stations are: Mendocino City, Mendocino County, and Michigan Bluff,
Placer County (Mus. Vert. Zool.) ; easternmost station at the south:
Campo, San Diego County (Mus. Vert. Zool.). Intergradation with
R. a. aurora takes place in western Mendocino and Humboldt counties.
Inhabits chiefly the Upper Sonoran life-zone, but extends into Trans-
ition and Lower Sonoran. Lives in damp places and about springs,
streams, and ponds. (See fig. 5.)
Rana pipiens Schreber
Leopard Frog
OriGInAL Description.—Rana pipiens Schreber, Naturforscher, 18,
1782, pp. 185-191.
Type Locauiry.—New York [State].
SynonymM.—Rana pipiens brachycephala.
Rance.—Along the Colorado River, north at least to near River-
side Mountain, Riverside County, and in the Imperial Valley, six
miles west of Imperial (Mus. Vert. Zool.). Life-zone, Lower Sonoran.
Inhabits river banks and fresh-water sloughs. (See fig. 5.)
Class REPTILIA
Order SQUAMATA
Suborder SAURIA
Family EUBLEPHARIDAE
Coleonyx variegatus (Baird)
Banded Gecko
Oriana Description.—Stenodactylus variegatus Baird, Proe.
Acad. Nat. Sci. Phila., 1858 [1859], p. 254.
150 University of California Publications in Zoology [Vou.17
Type Locaniry.—Rio Grande and Gila valleys: really Colorado
Desert, in southern California (see Stejneger, N. Amer. Fauna, 7,
1893, p. 163).
SynonyM.—Lublepharis variegatus.
Common Names.—Variegated Gecko; Variegated Lizard.
RancGE.—Southeastern desert areas. Occurs north to Big Pine,
Owens Valley, Inyo County (Mus. Vert. Zool.) ; west to Mohave, Kern
County (Stejneger, loc. cit., p. 164), to Colton, San Bernardino County
(Mus. Vert. Zool.), to San Jacinto, Riverside County (Van Denburgh,
Oce. Papers Calif. Acad. Sei., 5, 1897, p. 41), and to La Puerta Valley,
San Diego County (Stephens, MS.). Restricted to the Lower Sonoran
life-zone. Lives under stones and other objects and in burrows in
the ground. (See fig. 9.)
Family IGUANIDAE
Subfamily IguanrmaE
Dipso-saurus dorsalis (Baird and Girard)
Desert Iguana
Original DeEscription.—Crotaphytus dorsalis Baird and Girard,
Proce. Acad. Nat. Sei. Phila., 6, 1852, p. 126.
Typr Locauiry.—Desert of Colorado, California.
Common Names.—Crested Lizard; Colorado Desert Lizard; Keel-
backed Lizard; Thirsty Lizard; Sharp-back Lizard.
Rance.— Extreme desert areas east of the Sierran divides. Occurs
north to Owens, Panamint, Death (Mesquite), and Amargosa valleys
(Stejneger, N. Amer. Fauna, 7, 1893, p. 165) ; west to Coyote Wells,
Imperial County (Cope, Rep. U. S. Nat. Mus., 1898 [1900], p. 245) ;
Torres [Toro], Riverside County (Mus. Vert. Zool.) ; Daggett, San
Bernardino County (Meek, Field Columb. Mus., zool. ser., 7, 1905
[1906], p. 7), and 3 miles east of Owens Lake, Inyo County (Stejneger,
loc. cit.). Restricted to the Lower Sonoran life-zone. Lives on level,
sandy, or gravelly ground.
Uma notata Baird
Ocellated Sand Lizard
ORIGINAL Description.—Uma notata Baird, Proce. Acad. Nat. Sei.
Phila., 1858 [1859], p. 253.
1917] Grinnell-Camp: Amphibians and Reptiles of California 151
Type Locauiry.—Mohave Desert [== Colorado Desert, in Cali-
fornia ?].
Synonyms.—Uma inornata Cope, Amer. Nat., 29, 1895, p. 939
(type from Colorado Desert, ‘‘San Diego’’ [= Imperial] County) ;
Uma rufopunctata.
Common Names.—Ocellated Desert Lizard; Red-spotted Desert
Lizard; Cope’s Desert Lizard; Spotted Yuma Lizard.
RanGe.—Mohave and Colorado deserts; occurs west to near Signal
Mountain, Imperial County (Mus. Vert. Zool.), and north at least to
Daggett, San Bernardino County (Meek, Field Columb. Mus., zool.
ser., 7, 1906, pp. 4-7), and near Blythe Junction, in Riverside County
(Camp, Univ. Calif. Publ. Zool., 12, 1916, pp. 516-519). Oceurs only
within the Lower Sonoran life-zone, where further restricted to tracts
of eolian sand.
Callisaurus ventralis ventralis (Hallowell)
Southern Gridiron-tailed Lizard
ORIGINAL DEscripTION.—Homolosaurus [corrected in ‘‘ Errata’’ to
Homalosaurus| ventralis Hallowell, Proe. Acad. Nat. Sei. Phila., 6,
1852, pp. 179-180.
Type Locaurry.—New Mexico [west of the Rio Grande].
Synonyms.—Callisaurus draconoides ; Callisaurus draconoides ven-
tralis; Callisaurus dracontoides gabbu.
JomMON NamMeEs.—Zebra-tailed Lizard ; Spotted-tail Dragon ; Beau-
tiful Lizard.
Rance.—Entire desert area of southeastern California. Occurs
north into Inyo County: Independence in Owens Valley (Swarth,
MS), Saline Valley, Furnace Creek in Death Valley, and the Funeral
Range (Stejneger, N. Amer. Fauna, 7, 1893, p. 173); west to Kelso
Creek, near Weldon, Kern County (Mus. Vert. Zool.), to Cameron,
8 miles northwest of Mohave, Kern County (Stejneger, loc. cit.), to
western Antelope Valley, Los Angeles County (Camp, MS), to Cajon
Wash, San Bernardino County (Mus. Vert. Zool.), and to near Ban-
ning, Riverside County (Van Denburgh, Oce. Papers Calif. Acad. Sei.,
5, 1897, p. 50). Oceurs also at Vallevista, in San Jacinto Valley,
Riverside County (Atsatt, Univ. Calif. Publ. Zool., 12, 1913, p. 33),
and Oak Springs, San Diego County (Van Denburgh, loc. cit.). Oc-
cupies the Lower Sonoran lhfe-zone. Inhabits level gravelly or sandy
plains and washes.
152 University of California Publications in Zoology [Vou.17
Crotaphytus wislizenii Baird and Girard
Great Basin Leopard Lizard
OriGiInaL Description.—Crotaphytus Wislizenii Baird and Girard,
Proe. Acad. Nat. Sei. Phila., 6, 1852, p. 69.
Type Locauiry.—Near Santa Fé, New Mexico.
Synonyms.—Crotaphytus Gambelii Baird and Girard, loc. cit.,
p. 126 (type possibly from California: see Stejneger, N. Amer. Fauna,
3. 1890, p. 105, footnote) ; Crotaphytus fasciatus, part; [?] Crota-
phytus silus, part.
Common Nameb.—Wislizenius’ Lizard, part.
Ranee.—Arid interior chiefly east of the desert divides. Occurs
north through the Inyo region to Benton, Mono County (Mus. Vert.
Zool.) ; also recorded from Doyle, Lassen County (Richardson, Proe.
U.S. Nat. Mus., 48, 1915, p. 407). Oceurs west across Owens Valley
to Carroll Creek, west of Owens Lake, Inyo County (Mus. Vert. Zool.) ;
through Kern gap in vicinity of Walker Pass to Bodfish, Kern County
(Mus. Vert. Zool.) ; through Tejon Pass and Cuddy Canon to Lock-
wood Valley, 5000 feet altitude, Ventura County (Meek, Field Columb.
Mus., zool. ser., 7, 1905 [1906], p. 8); to 2 miles west of Pallett, Los
Angeles County (Mus. Vert. Zool.) ; to Cushenbury Springs, San
Bernardino County (Grinnell, Univ. Calif. Publ. Zool., 5, 1908, p.
160); through San Gorgonio Pass to Cabezon, Riverside County
(Atsatt, Univ. Calif. Publ. Zool., 12, 1913, p. 34) ; also to San Jacinto,
Riverside County (Van Denburgh, Oce. Papers Calif. Acad. Sci., 5,
1897, p. 58); to Pinyon Flat, 4000 feet altitude, on east slope Santa
Rosa Mountains, Riverside County (Atsatt, loc. cit.); and to Oak
Grove, San Diego County (MeLain, Crit. Notes Coll. Reptiles Western
U. S., 1899, p. 2). Oceupies the Lower and Upper Sonoran life-zones.
Inhabits level or rolling ground of sandy or gravelly nature.
Crotaphytus silus Stejneger
San Joaquin Leopard Lizard
ORIGINAL Description.—Crotaphytus silus Stejneger, N. Amer.
Fauna, 3, 1890, p. 105.
Typs Locauiry.—Fresno, Fresno County, California.
Synonyms.—Crotaphytus wislizenti, part; Crotaphytus fasciatus,
part.
Common Names.—Short-nosed Leopard Lizard; Wislizenius’ Liz-
ard, part; Banded Lizard.
1917} Grinnell-Camp: Amphibians and Reptiles of California 153
RanGE.—Southern San Joaquin Valley (Tulare basin), and Car-
rizo Plain, San Luis Obispo County. Occurs north to Livingston,
Merced County (MeLain, Crit. Notes Coll. Reptiles Western U. S.,
1899, p. 2); east to Poso, Bakersfield, and Pampa, in Kern County
(Stejneger, N. Amer. Fauna, 7, 1893, p. 170) ; south to 5 miles north
of Rose Station (Stejneger, loc. cit.), and to San Emigdio Plains
(Mus. Vert. Zool.), Kern County; then, on Carrizo Plain, west to
Painted Rock, San Luis Obispo County. Restricted to the Lower
Sonoran life-zone. Inhabits level valley floors.
Crotaphytus collaris baileyi Stejneger
Bailey Collared Lizard
OriginaL Description.—Crotaphytus baileyi Stejneger, N. Amer.
Fauna, 3, 1890, pp. 103-105, pl..12, fig. 1.
TypE Locaniry.—Painted Desert, Little Colorado River, Arizona.
Synonym.—Crotaphytus collaris.
Common Name.—Bailey Leopard Lizard.
RANGE.—Of interrupted distribution on the southeastern deserts,
occurring north through the Inyo region to Deep Spring Valley slope
of White Mountains, 5600 feet altitude, Inyo County (Stejneger, N.
Amer. Fauna, 7, 1893, p. 166). Has been taken west to 5 miles north
of Kernville, Kern County (Mus. Vert. Zool.) ; to Lytle Creek, San
Bernardino County (Van Denburgh, Proe. Calif. Acad. Sei., ser. 4,
3, 1912, p. 147); to Hall Grade, near Cabezon, Riverside County
(Atsatt, Univ. Calif. Publ. Zool., 12, 1913, p. 33); to Palm Canon,
San Jacinto Mountains, Riverside County (Atsatt, loc. cit.) ; and to
Mountain Spring, eastern San Diego County (Cope, Ann. Rep. U.S.
Nat. Mus., 1898 [1900], p. 252). Not known from region of Colorado
River south of near Blythe Junetion, in Riverside County (Camp,
Univ. Calif. Publ. Zool., 12, 1916, p. 521). Occurs in the Lower and
Upper Sonoran life-zones. Restricted to rocky situations.
Sauromalus ater Duméril
Chuckwalla
OriGInaAL DerscripTion.—Sauromalus ater Duméril, Arch. Maus.
Hist. Nat., 8, 1856, pp. 536-538, pl. 23, figs. 3, 3a.
Type Locauiry.—Unknown.
Synonym.—Luphryne obesus Baird, Proe. Acad. Nat. Sei. Phila.,
154 University of California Publications in Zoology [Vou.17
1858 [1859], p. 253 (type no. 4172, U. S. Nat. Mus., from Fort Yuma,
California).
Common Names.—Alderman Lizard; Fat Toad Lizard; Fat Lizard.
Rance.—Desert mountains of southeastern California. Occurs
north to Willow Creek, 4500 feet altitude, Panamint Mountains, Inyo
County (Stejneger, N. Amer. Fauna, 7, 1893, p. 175) ; west to Shep-
herd Canon, Argus Range, Inyo County (Stejneger, loc. cit.) ; to
Barstow and Victorville, San Bernardino County (Mus. Vert. Zool.) ;
to Snow Creek, San Jacinto Mountains, and Dos Palmos Spring, Santa
Rosa Mountains, in Riverside County (Atsatt, Univ. Calif. Publ. Zool.,
12, 1913, p. 34); and to Coyote Well, Imperial County (Cope, Ann.
Rep. U. S. Nat. Mus., 1898 [1900], p. 268), and east slope of ‘‘ Julian
Mountains’’ [== Cuyamaca Mountains?], in eastern San Diego County
(Van Denburgh, Oce. Papers Calif. Acad. Sei., 5, 1897, p. 62). Re-
strieted to the Lower Sonoran life-zone, where it occurs only in rocky
situations.
Uta mearnsi Stejneger
Mearns Lizard
OriegInaL Description.—Uta mearnsi Stejneger, Proe. U. 8. Nat.
Mus., 17, November, 1894, pp. 589-591.
Type Locauiry.—Summit of Coast Range, United States and
Mexican boundary line, California.
Common Name.—Mearns Swift.
Rance.—Desert slopes of San Jacinto, Santa Rosa and Coast
ranges, In Riverside and San Diego counties. Has been taken north-
east to Banning, Cabezon, Snow Creek and Palm Canon, in Riverside
County (Atsatt, Univ. Calif. Publ. Zool., 12, 1913, pp. 34-35), west
to La Puerta, San Diego County (Mus. Vert. Zool.), and south to the
type locality, as above given. Altitudes of occurrence extend from
800 to 3500 feet. Oceurs in the Lower Sonoran life-zone, extending
loeally into Upper Sonoran. Lives on boulders and faces of cliffs.
Uta stansburiana stansburiana Baird and Girard
Northern Brown-shouldered Lizard
OrIGINnAL DescripTion.—Uta Stansburiana Baird and Girard, Proe.
Acad. Nat. Sei. Phila., 6, 1852, p. 69.
Type Locaniry.—Valley of Great Salt Lake, Utah.
Ranee.—Enters California from Nevada in the Inyo region.
1917] Grinnell-Camp: Amphibians and Reptiles of California — 155
Specimens recorded from Round Valley and near Lone Pine, in Inyo
County (Richardson, Proc. U. S. Nat. Mus., 48, 1915, p. 412). Addi-
tional specimens in Mus. Vert. Zool., from vicinity of Independence
and Laws, Inyo County, and from Benton, Mono County. Oceupies
Uta stansburiana stansburiana
Intergrade
Uta stansburiana elegans
Intergrade 4
Uta stansburiana hesperis
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA
Fig. 6. Distribution of Uta stansburiana in California.
the Upper Sonoran life-zone. Inhabits chiefly sandy or gravelly
ground among bushes. (See fig. 6.)
Uta stansburiana elegans Yarrow
Desert Brown-shouldered Lizard
OrigInaL Description.—Uta elegans Yarrow, Proce. U. 8. Nat.
Mus., 5, 1882, p. 442.
156 Unwersity of California Publications in Zoology [Vou.17
Tyre Locauiry.—lLa Paz, Lower California, Mexico.
Synonym.—Uta stansburiana, part.
Common Names.—Stansbury’s Swift, part ; Stansbury’s Uta, part ;
Brown-shouldered Lizard, part.
Rancre.—tThe southeastern deserts. Has been captured north to
Carroll Creek and Keeler, Inyo County; west to the Pacific divides,
where intergradation takes place with U. s. hesperis. Intergradation
with U. s. stansburiana takes place in Owens Valley, Inyo County
(Mus. Vert. Zool.). Oceupies chiefly the Lower Sonoran life-zone.
Inhabits nearly all types of desert environment except sand dunes.
(See fig. 6.)
Uta stansburiana hesperis Richardson
Southern Brown-shouldered Lizard
OriginaL Derscription.—Uta stansburiana hesperis Richardson,
Proc. U. S. Nat. Mus., 48, January 19, 1915, pp. 415-418.
Type Locaniry.—Arroyo Seeo Canon, near Pasadena, Los Angeles
County, California.
SynonyM.—Uta stansburiana, part.
Common Names.—Brown-shouldered Lizard, part; Stansbury’s
Swift, part; Stansbury’s Uta, part.
Rance.—Coastal slopes of southern California and the San Joa-
quin Valley. Northernmost stations, Mount Diablo, Contra Costa
County (Univ. Calif. Dept. Zool.), and 5 miles south of Lathrop, San
Joaquin County (Van Denburgh, Oee. Papers Calif. Acad. Sei., 5,
1897, p. 68; Camp, Univ. Calif. Publ. Zool., 17, 1916, p. 69). Extends
northeast to Fresno and west in central California to Bear Valley,
San Benito County (Van Denburgh, loc. cit.). Occurs also on Santa
Cruz, Ana Capa, Santa Catalina, and San Clemente islands. Inter-
grades on the east, through Walker Pass and across the desert divides
in southern California, with U. s. elegans (Mus. Vert. Zool.). Oc-
cupies the Lower Sonoran life-zone and to a lesser extent the Upper
Sonoran life-zone. Inhabits arid plains, washes, the more open, dry
hill-slopes, and, along the seacoast, sand dunes. (See fig. 6.)
Uta graciosa (Hallowell)
Long-tailed Uta
OriginaL Description.—Uro-saurus graciosus Hallowell, Proce.
Acad. Nat. Sci. Phila., 7, 1854, pp. 92-93.
ol
1917} Grinnell-Camp: Amphibians and Reptiles of California 1
Type Locaniry.—Lower [=southern?] California.
SynonyM.—Anolis coopert Baird, Proe. Acad. Nat. Sei. Phila.,
1858 [1859], p. 254 (type from California).
Common Namus.—Long-tailed Swift; Graceful Uta; Cooper’s
Green Lizard; Slender Lizard.
RancGe.—Colorado Desert, in vicinity of Colorado River, and west
at least to Blythe Junction and Goffs (Blake P. O.), Riverside and
San Bernardino counties (Camp, Univ. Calif. Publ. Zool., 12, 1916,
pp. 525-526), and Meloland, Imperial County (Mus. Vert. Zool.).
Restricted to the Lower Sonoran life-zone. Lives in bushes and desert
trees.
Uta ornata Baird and Girard
Tree Uta
OrIGINAL Description.—Uta ornata Baird and Girard, Proce. Acad.
Nat. Sci. Phila., 6, 1852, p. 126.
Type Locauiry.—Rio San Pedro (Texas) and province of Sonora
[= Arizona].
Synonyms.—Uta symmetrica Baird, Proce. Acad. Nat. Sei. Phila.,
1858 [1859], p. 253 (type from Fort Yuma) ; [?] Anolis carolinensis ;
[?] Anolis principalis; [2] Uta schottii Baird, loc. cit. (type from
“Sta. Madelina, Cal.’’).
Common Names.—Ornate Swift ; White-bellied Swift ; Ornate Uta;
Tree Swift; Graceful Lizard; [?] Schott’s Lizard.
Rance.—Valley of the Colorado River north at least to vicinity
of Palo Verde, Imperial County (Mus. Vert. Zool.). Restricted to
the Lower Sonoran life-zone. Lives on trunks of trees in the river
bottom.
Sceloporus graciosus graciosus Baird and Girard
Mountain Lizard
Ori@inaL Description.—Sceloporus graciosus Baird and Girard,
Proe. Acad. Nat. Sei. Phila., 6, 1852, p. 69.
Type Locauiry.—Valley of the Great Salt Lake [Utah].
Synonyms.—|[?] Sceloporus consobrinus; Scelopurus consobrinus
gratiosus.
Common Names.—New Mexican Alligator Lizard; Sage-brush
Swift, part; Fence Lizard, part; Marey’s Alligator Lizard.
RaNGE.—Chiefly mountainous districts from 2400 feet altitude in
the southern Sierras (Bodfish, Kern County) and 700 feet in the inner
158 University of California Publications in Zoology (Vou. 17
northern coast ranges (3 miles west of Vacaville, Solano County) to
8100 feet in Mariposa County (near Porcupine Flat) (Mus. Vert.
Zool.). Oceurs typically south to Mount Pinos, Ventura County.
Ranges west to South Yolla Bolly Mountain, 6 miles south of Covelo,
Sceloporus graciosus graciosus
Intergrade
Sceloporus graciosus vandenburgianus
Sceloporus magister
Lbeeond
Sceloporus orcutti
Range of Sceloporus graciosus outlined
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA,
Fig. 7. Distribution of Sceloporus graciosus, S. magister, and S. orcutti in
California.
and Mount Sanhedrin, in Trinity and Mendocino counties (Mus. Vert.
Zool.) ; and south in the Sierra Nevada as far as Mount Breckinridge,
6500 feet altitude, in Kern County (Mus. Vert. Zool.). Occurs also
on the Warner Mountains, from 4700 to 5000 feet altitude, Modoe
County (Mus. Vert. Zool.) ; about Mono Lake and at Benton, in Mono
County (Mus. Vert. Zool), and on the Panamint Mountains, 6400 feet
1917] Grinnell-Camp: Amphibians and Reptiles of California 159
altitude, Inyo County (Stejneger, N. Amer. Fauna, 7, 1893, p. 184).
Oceupies chiefly the Transition life-zone, but extends locally into the
Canadian on the one hand and more commonly into the Upper Sonoran
on the other. Lives on and about rocks and logs and on the ground
among bushes. (See fig. 7.)
Sceloporus graciosus vandenburgianus Cope
Van Denburgh Lizard
OriGinaL Drscriprion.—NSceloporus vandenburgianus Cope, Amer.
Nat., 30, 1896, pp. 834-836.
Type Locaurry.—Summit of Coast Range [probably Laguna
Mountains], San Diego County, California.
SYNONYM.
Common NAmes.—Mountain Lizard, part; Van Denburgh’s Swift ;
Sage-brush Swift, part.
RancGe.—Cuyamaea, Santa Rosa, San Jacinto, San Bernardino, and
San Gabriel mountains, from 4000 to 8500 feet altitude. Northwest-
ernmost station: Pine Flats, in San Gabriel Mountains, Los Angeles
Sceloporus graciosus, part.
County (Mus. Vert. Zool.). Intergradation with S. g. graciosus is
shown by specimens from San Bernardino and San Gabriel mountains
(see Camp, Univ. Calif. Publ. Zool., 17, 1916, pp. 67-68). Oceupies
the Transition hfe-zone. Lives on and about rocks and logs and on
the ground among bushes. (See fig. 7.)
Sceloporus occidentalis occidentalis Baird and Girard
Pacific Blue-bellied Lizard
OrIGINAL DescripTion.—Sceloporus occidentalis Baird and Girard,
Proc. Acad. Nat. Sci. Phila., 6, 1852, p. 175.
Type Locauiry.—California [by inference from next subsequent
published definite statement, Benicia (Baird, Pace. R. R. Rep., 10, 1859,
Abbot’s Rep., pt. 4, no. 4, p. 9) ].
Synonyms.—Sceloporus undulatus var. bocourtii Boulenger, Cat.
Lizards Brit. Mus., ed. 2, 2, 1885, p. 229, part (specimens listed from
Monterey, Mt. Whitney, and Santa Cruz [!]); Sceloporus frontalis;
Sceloporus undulatus occidentalis, part; Sceloporus undulatus undu-
latus, part; Sceloporus undulatus thayert, part.
Common Names.—Western Fence Lizard; Western Alligator
Lizard, part; Pacific Swift; Thayer’s Alhigator Lizard, part ; Alligator
Lizard, part.
160 University of California Publications in Zoology |Vou.17
RaNncGE.—Northern and west-central portions of the state, including
both the coast district and Sacramento and San Joaquin valleys.
Occurs east to the Warner Mountains, Modoe County, and into the
foothills of the Sierra Nevada as far as Fyffe, Eldorado County, and
Yosemite Valley, Mariposa County (Mus. Vert. Zool.) ; south to Snell-
ing and 22 miles south of Los Banos, in Merced County, and to San
Luis Obispo (Mus. Vert. Zool.). In the vicinity of the latter points
intergradation with S. o. bi-seriatus takes place, as also in eastern
Modoe County (see Camp, Univ. Calif. Publ. Zool., 17, 1916, pp. 63-
65). Oceupies the Upper Sonoran and Transition life-zones. Lives
on tree-trunks, fences, logs, boulders, and in steep banks. (See fig. 8.)
Sceloporus occidentalis taylori Camp
Tenaya Blue-bellied Lizard
OriGINaL Derscription.—Sceloporus - occidentalis taylori Camp,
Uniy. Calif. Publ. Zool., 17, December 28, 1916, pp. 65-67.
Type Locaurry.—Half-way between Merced Lake and Sunrise
Trail, altitude 7500 feet, Yosemite National Park, Mariposa County,
California.
Rance.—Upper basins of Tuolumne and Merced rivers, between
altitudes of 7300 and 8200 feet, in Yosemite National Park. Occupies
the Canadian life-zone. Lives on and beneath boulders and in rock-
slides. (See fig. 8.)
Sceloporus occidentalis bi-seriatus Hallowell
Fence Lizard
ORIGINAL DeEscriptionN.—Sceloporus bi-seriatus Hallowell, Proce.
Acad. Nat. Sei. Phila., 7, 1854, pp. 93-94.
Type Locauiry.—Borders of El Paso Creek and in Tejon Valley
[both in Kern County, California}.
Synonyms.—Sceloporus longipes Baird, Proe. Acad. Nat. Sci.
Phila., 1858 [1859], p. 254 (type from Fort Tejon, California) ;
Sceloporus undulatus bocourtii, part [see under S. 0. occidentalis] ;
Sceloporus undulatus thayeri, part; Sceloporus undulatus undulatus,
part; Sceloporus [bi-seriatus] var. marmoratus; Sceloporus occiden-
talis, part; Sceloporus bi-seriatus var. A. azwreus Hallowell, loc. cit.,
p. 94 (type locality same as for S. bi-sertatus) ; Sceloporus bi-seriatus
var. B. variegatus Hallowell, loc. cit., p. 94 (type locality same as for
S. bi-seriatus).
1917] Grinnell-Camp: Amphibians and Reptiles of California 161
Common Names.—Fence Swift; Western Swift ; Two-lined Lizard ;
Thayer’s Alligator Lizard, part; Western Alligator Lizard, part;
Blue-bellied Lizard, part; Common Swift; Long-footed Lizard ; Two-
striped Lizard.
Sceloporus occidentalis occidentalis
Intergrade |
Sceloporus occidentalis bi-seriatus
Intergrade
Sceloporus occidentalis becki 4
71o8 gd e080
Sceloporus
occidentalis taylori
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OP CALIFORNIA
a
Fig. 8. Distribution of Blue-bellied Lizards (Sceloporus occidentalis) in
California.
Rance.—East-central and southwestern portions of the state.
Occurs north to Matilija, Ventura County, to Carrizo Plain, San Luis
Obispo County, to Raymond, Madera County, and to Mono Lake,
Mono County (Mus. Vert. Zool.) ; east to Benton, Mono County, to
Carroll Creek, Inyo County, to west slope of Walker Pass, Kern
162 University of California Publications in Zoology [Vou.17
County, to Pallett, Los Angeles County, to Victorville and Cushen-
berry Springs, San Bernardino County, to Cabezon, Strawberry Valley,
and Santa Rosa Mountains at 6000 feet altitude, in Riverside County,
and to Warner Pass and Jacumba (2825 feet altitude), in eastern
San Diego County (all above localities represented by specimens in
Mus. Vert. Zool.). Extends altitudinally as high as 10,000 feet near
Kearsarge Pass on eastern declivity of Sierra Nevada in Inyo County
(Mus. Vert. Zool.). Occurs also, in the Inyo region, on the Coso,
Argus, Panamint, Inyo, and White mountains (Stejneger, N. Amer.
Fauna, 7, 1893, pp. 185-186). Oceupies the Lower and Upper Sonoran
life-zones and extends locally into Transition. Lives on tree-trunks,
fences, sides of buildings, and among rocks of large size. (See fig. 8.)
Sceloporus occidentalis becki Van Denburgh
Island Blue-bellied Lizard
ORIGINAL DEscRIPTION.—Sceloporus becki Van Denburgh, Proce.
Calif. Acad. Sci., ser. 3, zool., 4, June 15, 1905, pp. 9-10, pl. 4.
Type Locauiry.—San Miguel Island, Santa Barbara County,
California.
Synonym.—Sceloporus biseriatus becki.
RanceE.—The extreme form of this subspecies occurs only at the
type locality ; forms intermediate between this and the mainland sub-
species are found, according to Van Denburgh (loc. cit.) on Santa
Rosa and Santa Cruz islands, Santa Barbara County. Lives among
large rocks and on tree-trunks. (See fig. 8.)
Sceloporus magister Hallowell
Desert Rough-sealed Lizard
ORIGINAL Description.—Sceloporus magister Hallowell, Proc.
Acad. Nat. Sei. Phila., 7, 1854, p. 93.
Type Locauiry.—Near Fort Yuma, California.
Synonyms.—Sceloporus clarkii; Sceloporus spinosus var. clarki;
[2] Sceloporus orcutts, part.
Common Names.—Sealy Lizard; Clark’s Alligator Lizard; Fence
Lizard, part; Clark’s Lizard; Great Fence Lizard.
Raneu.—Entire southeastern desert area, extending north to Lone
Pine (Stejneger, N. Amer. Fauna, 7, 1893, p. 183), to Mazourka Canon,
Inyo Mountains (Mus. Vert. Zool.), and to Willow Creek, Panamint
1917] Grinnell-Camp: Amphibians and Reptiles of California 163
Mountains (Stejneger, loc. cit.), im Inyo County; west through the
Kern River gap to Bodfish, Kern County (Mus. Vert. Zool.) ; also
west to Fairmont and Pallett, Los Angeles County (Mus. Vert. Zool.) ;
to Hesperia (Van Denburgh, Oce. Papers Calif. Acad. Sci., 5, 1897,
p. 86) and Cushenbury Springs (Grinnell, Univ. Calif. Publ. Zool.,
5, 1908, p. 162), in San Bernardino County; to Cabezon and Dos
Palmos Spring (38000 feet altitude), in Riverside County (Atsatt,
Univ. Calif. Publ. Zool., 12, 1913, p. 37); and to Warner Pass and
La Puerta, San Diego County (Mus. Vert. Zool.). There is one record
for the west side of the San Joaquin Valley: Los Gatos Cafion, 6 miles
north of Coalinga, in Fresno County (Van Denburgh, loc. cit., p. 86).
Occupies the Lower Sonoran life-zone. Inhabits tree-yuceas, catclaw
thickets, piles of rocks, undercut wash-banks, and railroad culverts.
(See fig. 7.)
Sceloporus orcutti Stejneger
Dusky Rough-sealed Lizard
ORIGINAL DescripTion.—Sceloporus orcutti Stejneger, N. Amer.
Fauna, 7, May, 1893, p. 181 (footnote), pl. 1, figs. 4a-4c.
Type Locaurry.—Milquatay Valley [about 50 miles east of San
Diego], San Diego County, California.
Synonym.—[?] Sceloporus spinosus.
Common Names.—Oreutt’s Swift; Dusky Sealy Lizard; [?] Spiny
Alligator Lizard.
Rance.—Mountains of San Diego, western Riverside, and south-
western San Bernardino counties. Occurs northwest to Waterman
Canon, San Bernardino County (Van Denburgh, Proe. Calif. Acad.
Sei., ser. 4, 3, 1912, p. 149) ; west to Riverside (Van Denburgh, loc.
cit.) and Temeseal (Van Denburgh, Oce. Papers Calif. Acad. Sei., 5,
1897, p. 88), in Riverside County, and to Escondido, San Diego County
(Mus. Vert. Zool.) ; east to Mountain Spring, San Diego County (Cope,
Ann. Rep. U. 8. Nat. Mus., 1898 [1900], p. 356) ; and to Dos Palmos
Spring (3500 feet altitude), Snow Creek (2000 feet), and Banning
(2200 feet), in Riverside County (Atsatt, Univ. Calif. Publ. Zool., 12,
1913, p. 37). Extends as high as 5900 feet altitude on west side of San
Jacinto Mountains, at Fuller’s Mill (Atsatt, loc. cit.). Oceurs chiefly
within the Upper Sonoran life-zone. Lives among boulders surrounded
by chaparral. (See fig. 7.)
164 University of California Publications in Zoology [Vou.17
Phrynosoma douglassii dougilassii (Bell)
Pigmy Horned-toad
OriciInAL Description.—Agama Douglassii Bell, Trans. Linn. Soe.
Lond., 16, 1828, pp. 105-107, pl. 10.
Type Locaniry.—Columbia River.
Synonyms.—Phrynosoma douglassi pygmaea; Tapaya Douglassu.
Common Names.—Pigmy Horned Lizard ; Douglass’s Horned Toad.
Ranee.—One definite record from the state: western base of Mount
Shasta [= Shasta Valley?], Siskiyou County (Townsend, Proce. U.S.
Nat. Mus., 10, 1887, p. 238).
Phrynosoma, blainvillii blainvillii (Gray)
Blainville Horned-toad
OriGgInaAL Description.—Phrynosoma Blainvillii Gray, Zool.
Beechey’s Voyage, 1839, p. 96, pl. 29, fig. 1.
Typr Locaniry.—California: probably San Diego (see Van Den-
burgh, Proce. Calif. Acad. Sci., ser. 2, 4, 1894, p. 296).
Synonyms.—Agama coronatum; Phrynosoma coronatum, part;
Phrynosoma modestum.
Common Namus.—Pacifie Horned Lizard; Blainville’s Horned
Lizard, part; California Horned Lizard, part; Pacifie Horned Toad ;
Crowned Horned Lizard, part; Little Horned Lizard.
RANnGE.—Mainly west of the desert divides, in San Diego, Orange,
Riverside, San Bernardino, and Los Angeles counties. Occurs north
to Tujunga Wash, near Sunland, Los Angeles County, and to Cajon
Pass, San Bernardino County (Van Denburgh, Proe. Calif. Acad. Sci.,
ser. 4, 3, 1912, p. 148) ; east to Jacumba (Cope, Ann. Rep. U.S. Nat.
Mus., 1898 [1900], p. 427), and Warner Pass (Bryant, Univ. Calif.
Publ. Zool., 9, 1911, p. 36), m San Diego County; to Vandeventer
Flat, Oak Springs (west side of Palm Canon, 6500 feet altitude), and
Cabezon, in Riverside County (Bryant, loc. cit.) ; and to junction of
South Fork (6200 feet altitude) and upper Santa Ana Canon, San
Bernardino Mountains (Grinnell, Univ. Calif. Publ. Zool., 5, 1908,
p. 162). Oceupies the Upper and Lower Sonoran life-zones. Inhabits
eravelly or sandy ground of a nature to permit burrowing.
1917] Grinnell-Camp: Amphibians and Reptiles of California 165
Phrynosoma blainvillii frontale Van Denburgh
California Horned-toad
OrigINnAL Description.—Phrynosoma frontalis Van Denburgh,
Proce. Calif. Acad. Sci., ser. 2, 4, July 12, 1894, p. 296.
Typp Locauiry.—Bear Valley, San Benito County, California.
Synonyms.—Phrynosoma blainvilla, part ; Phrynosoma coronatum,
part; Phrynosoma cornutum; Batrachosoma coronatum; Tapaya
coronata.
Common Names.—California Horned Lizard, part ; Spiny-breasted
Horned Lizard; Blainyille’s Horned Lizard, part; Crowned Horned
Lizard, part.
Rance.— West-central California, chiefly west of the desert divides.
Oceurs north along east side of the Sacramento Valley at least to
Colfax, Placer County, and along west side of the San Joaquin
Valley to Traey, San Joaquin County (Bryant, Univ. Calif. Publ.
Zool., 9, 1911, p. 42) ; west to Berkeley, Alameda County (Mus. Vert.
Zool.), and to Searsville, San Mateo County, and Pacific Grove,
Monterey County (Van Denburgh, Oce. Papers Calif. Acad. Sci., 5,
1897, p. 95) ; east to five miles northeast Coulterville (3100 feet alti-
tude), in Mariposa County (Mus. Vert. Zool.), and Walker Pass (5200
feet altitude), in Kern County (Bryant, loc. cit.) ; southeast to 5 miles
south of Neenach, at 4000 feet altitude, and upper Tujunga Canton,
4500 feet altitude, in northern Los Angeles County (Mus. Vert. Zool.).
Intergrades south of latter points, between San Francisquito Canon
and Pasadena, with P. b. blainvillu. Inhabits the Upper and Lower
Sonoran life-zones. Lives in arid brushy or open situations.
Phrynosoma platyrhinos Girard
Desert Horned-toad
OrieInAL Description.—Phrynosoma platyrhinos Girard, Stans-
bury’s Expl. Gt. Salt Lake, 1853, pp. 361, 363-864, pl. 7, figs. 1-5.
Type Locaniry.—Great Salt Lake, Utah.
Synonyms.—Anota calidiarum Cope, Amer. Nat., 30, 1896, pp.
833-834 (type from Death Valley [uncertain]); Phrynosoma cali-
diarum; Dolrosaurus platyrhinos; Anota platyrhina; Phrynosoma
coronatum, part.
Common Names.—Desert Horned Lizard; Ashy Horned Toad;
166 University of California Publications in Zoology [Vou.17
Smooth Horned Lizard; Smooth Horned Toad; Broad-nosed Barrel
Lizard; Broad-nosed Horned Toad.
Rance.—The southeastern deserts chiefly east of the Pacific divides.
Recorded north through the Inyo region to Benton, Mono County
(Mus. Vert. Zool.) ; west to Carroll Creek, Inyo County (Mus. Vert.
Zool.), to South Fork Kern River near Chimney Creek (3100 feet
altitude) and head of Kelso Valley (5300 feet altitude), in Kern
County (Bryant, Univ. Calif. Publ. Zool., 9, 1911, p. 49) ; to Fairmont,
in northern Los Angeles County (Grinnell and Grinnell, Throop Inst.
Bull., 35, 1907, p. 57) ; to Victorville, San Bernardino County (Mus.
Vert. Zool.) ; to Whitewater (Atsatt, Univ. Calif. Publ. Zool., 12, 1913,
p. 38), Palm Springs, and Mecea (Bryant, loc. cit.), in Riverside
County ; and to Coyote Wells, in Imperial County (Cope, Ann. Rep.
U.S. Nat. Mus., 1898 [1900], p. 447). Oceupies the Lower Sonoran
life-zone. Lives in open gravelly or sandy situations.
Phrynosoma m’callii (Hallowell)
Flat-tailed Horned-toad
OriGiInaL Description—Anota M’Callii Hallowell, Proce. Acad.
Nat. Sei. Phila., 6, 1852, p. 182.
Type Locauiry.—Great Desert of the Colorado, between Vallecito
and Camp Yuma, about 160 miles east of San Diego.
Synonym.—Doliosaurus me’calli.
Common Names.—Flat-tailed Horned Lizard; MeCall’s Horned
Lizard; MacCall’s Horned Toad.
Rance.—The Salton Basin, in Imperial and Riverside counties.
Recorded from: Mecea, Riverside County (Bryant, Univ. Calif. Publ.
Zool., 9, 1911, p. 59); Coyote Well (Bryant, loc. cit.), Salton Sea
(south end) (Bryant, loc. cit.), Fort Yuma (Cope, Ann. Rep. U. S.
Nat. Mus., 1898 [1900], p. 450), and the type locality (as above), in
Imperial County. Restricted to the Lower Sonoran life-zone. In-
habits open tracts of sandy alluvium.
Family ANGUIDAE
Gerrhonotus scincicauda scincicauda (Skilton)
California Alligator Lizard
ORIGINAL Description.—Tropidolepis scincicauda Skilton, Amer.
Journ. Arts. Sei., ser. 2, 7, 1849, p. 202, pl. opp. p. 464, figs. 1-3.
1917] Grinnell-Camp: Amphibians and Reptiles of California 167
Type Locauiry.—Dalles of the Columbia [Oregon].
SynonymMs.—Elgaria scincicauda; Gerrhonotus multicarinatus,
part.
Common Names.—Skink-tailed Lizard, part; Many-keeled Lizard,
part; Many-ribbed Lizard.
Gerrhonotus scincicauda scincicauda 1
Gerrhonotus scincicauda webbii
Gerrhonotus palmeri
Gerrhonotus coeruleus
Coleonyx variegatus
Testudo agassizti
Ranges of Gerrhonotus scincicauda and |
Gerrhonotus coeruleus outlined
eae |
4
¢-—- —----
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY A
Fig. 9. Distribution of Alligator Lizards (Gerrhonotus), Gecko (Coleonyx),
and Desert Tortoise (Testudo agassizii) in California.
Rance.—West-central portion of the state. Extends south near
the seacoast from Monte Rio, Sonoma County (Mus. Vert. Zool.) at
least to Pacifie Grove, Monterey County (Ven Denburgh, Oce. Papers
Calif. Acad. Sci., 5, 1897, p. 106). Extends north interiorly to Fair-
banks (Van Denburgh, loc. cit.) and 3 miles west of Covelo (Mus.
168 University of California Publications in Zoology [Vou.17
Vert. Zool.), in Mendocino County, and to Squaw Creek, Siskiyou
County (U. S. Nat. Mus.) ; southeast to Riverton, Eldorado County
(Van Denburgh, loc. cit.). Supposed to intergrade on the southeast
with G. s. webbu. Occurs also on San Miguel, Santa Rosa, and Santa
Cruz islands (Van Denburgh, Proe. Calif. Acad. Sci., ser. 3, zool., 4,
1905, pp. 3, 10-11, 12-13, 14). Occupies the Upper Sonoran life-zone,
extending locally into Transition. Inhabits the chaparral. (See fig. 9.)
Gerrhonotus scincicauda webbii Baird
San Diego Alligator Lizard
OrIGINAL DescripTION.—Gerrhonotus webbii Baird, Proce. Acad.
Nat. Sci. Phila., 1858 [1859], p. 255.
Type Locauiry.—Near San Diego, California.
Synonyms.—Gerrhonotus scincicauda ignavus; Gerrhonotus scin-
cicauda, part; Gerrhonotus multicarinatus, part.
Common Names.—Many-keeled Lizard, part; Webb’s Lizard.
RaANGE.—Southern California, chiefly west of the desert divides and
north along the lower west slopes of the Sierra Nevada as far as 3
miles northeast of Coulterville (3000 feet altitude), Mariposa County
(Mus. Vert. Zool.) ; extends northwest at least to Matilija, Ventura
County (Mus. Vert. Zool.). Easternmost stations: Jacumba, San
Diego County (Cope, Ann. Rep. U.S. Nat. Mus., 1898 [1900], p. 525) ;
Cabezon, Riverside County (Atsatt, Univ. Calif. Publ. Zool., 12, 1913,
pp. 88-389) ; Swartout Canon, San Bernardino County (Van Denburgh,
Proe. Calif. Acad. Sci., ser. 4, 3, 1912, p. 148) ; and Onyx, Kern County
(Mus. Vert. Zool.). Occurs also on Santa Catalina Island (Van
Denburgh, Proe. Calif. Acad. Sci., ser. 4, 4, 1914, pp. 182, 138). Oc-
cupies the Upper and Lower Sonoran life-zones, extending locally into
Transition. Inhabits the chaparral; also occurs about buildings. (See
fig. 9.)
Gerrhonotus palmeri Stejneger
Sierran Alligator Lizard
OrIGINAL Derscrietion.—Gerrhonotus scincicauda palmeri Stej-
neger, N. Amer. Fauna, 7, May 31, 1893, pp. 196-197.
Type Locauiry.—South Fork Kings River [in Fresno County],
California.
1917] Grinnell-Camp: Amphibians and Reptiles of California 169
SynonyM.—Gerrhonotus multicarinatus palmerit.
Common Name.—Mountain Alligator Lizard.
Rance.—Middle slopes of Sierra Nevada, from Tahoe City (Rich-
ardson, Proce. U. S. Nat. Mus., 48, 1915, pp. 424-425) south to Jackass
Meadow, 7750 feet altitude, Tulare County (Mus. Vert. Zool.). High-
est altitude of capture, about 8800 feet (Stejneger, loc. cit.). Occurs
also on Mount Breckenridge, 6500 feet altitude, Kern County (Mus.
Vert. Zool.). Occupies the Canadian life-zone, extending into the
Transition. Inhabits chaparral and underbrush on forest floors. (See
fig. 9.)
Gerrhonotus coeruleus Wiegmann
Coast Alligator Lizard
OrIGINAL DescripTion.—Gerrhonotus coeruleus Wiegmann, Oken’s
Isis, 1828, 21, pts. 3-4, p. 379.
Type Locanuiry.—Brazil [= probably San Francisco: see Peters,
in Duméril Boeourt, and Moquard, in Miss. Sei. au Mex., Rechereh.
Zool., pt. 3, sec. 1, 6° Livr., 1879, p. 355, and Stejneger, Proc. Biol.
Soe. Wash., 15, 1902, p. 37].
Synonyms.—Gerrhonotus Burnettii Gray, in Griffith’s Animal
King., 9, 1831, Syn. Reptilia, p. 64 (from America [= California] ) ;
Elgaria formosa Baird and Girard, Proe. Acad. Nat. Sci. Phila., 6,
1852, p. 175 (type from California) ; Gerrhonotus grandis; Gerrho-
notus multicarinatus, part; Gerrhonotus scincicaudus, part.
Common Names.—Burnett’s Alligator Lizard ; Skink-tailed Lizard,
part; Burnett’s Keeled Lizard; Green-brown Lizard; Many-keeled
Lizard, part.
RaNGE.—A narrow ecoastwise strip, from Cuddeback, Humboldt
County (Mus. Vert. Zool.) south to Pacifie Grove, Monterey County
(Van Denburgh, Occ. Papers Calif. Acad. Sci., 5, 1897, p. 110). Oc-
curs northeast to South Fork Salmon River, Siskiyou County (Mus.
Vert. Zool.). Easternmost stations: 4 miles south of South Yolla
Bolly Mountain, Tehama County; Lierly’s, near Mount Sanhedrin, in
Mendocino County; and Redwood Canon, near Oakland, Alameda
County (Mus. Vert. Zool.). Oceupies the Transition life-zone. In-
habits chaparral and openings in forests; found frequently along
streams. (See fig. 9.)
170 University of California Publications in Zoology [Vou.17
Family ANNIELLIDAE
Anniella pulchra pulchra Gray
Silvery Footless Lizard
OrigINAL DescripTion.—Anniella pulchra Gray, Ann. Mag. Nat.
Hist., ser. 2, 10, 1852, p. 440.
Type Locaurry.—California.
Synonym.—Anniella texana.
Common Names.—Blue Worm-snake, part; Blind Worm; Worm
Snake, part; Worm Lizard.
Rance— Chiefly southern coast district. Recorded north to Contra
Costa County (Van Denburgh, Proc. Calif. Acad. Sei., ser. 3, zool.,
4, 1905, p. 48), and San Ardo, Monterey County (Van Denburgh,
Oce. Papers Calif. Acad. Sci., 5, 1897, pp. 117-118) ; east to Bear
Valley, San Benito County (Van Denburgh, 1897, loc. cit.), Sequoia
National Park, Tulare County, and between Oil City and Poso Creek,
in Kern County (Van Denburgh, 1905, loc. cit.) ; to La Canada, near
Pasadena (Grinnell and Grinnell, Throop Inst. Bull., 35, 1907, p. 33),
San Bernardino and San Jacinto (Van Denburgh, 1897, loc. cit.), and
La Puerta Valley, in eastern San Diego County (Mus. Vert. Zool.).
Occurs within the Lower and Upper Sonoran life-zones. Inhabits
sandy ground; lives beneath rocks in dry washes and in sand dunes.
Anniella pulchra nigra Fischer
Black Footless Lizard
ORIGINAL DescripTioN.—Anniella nigra Fischer, Abh. Nat. Verein
Hamburg, 9, pt. 1, 1886, pp. 9-10, 1 pl.
Typr Locauiry—San Diego, California [more likely near Monte-
rey; see Van Denburgh, Proce. Calif. Acad. Sci., ser. 3, zool., 4, 1905,
p. 45].
Synonym.—Anniella pulchra, part.
Common Name.—Blue Worm Snake, part.
Rance.—Central seaboard. Recorded only from San Francisco
(Cope, Ann. Rep. U. S. Nat. Mus., 1898 [1900], pp. 675, 676), vicinity
of Monterey: Pacific Grove, Point Pinos, and Carmel Bay (Van
Denburgh, loc. cit., p. 48), and Marin County (Rivers, Bull. South.
Calif. Aead. Sci., 1, March, 1902, p. 27). Specimens examined by us
(in Coll. San Diego Soe. Nat. Hist.) from Morro Bay, San Luis Obispo
1917] Grinnell-Camp: Amphibians and Reptiles of California 171
County, are intermediate towards pulchra, but nearest nigra. Occurs
within the Transition life-zone. Lives in sand dunes.
Family XANTUSIIDAE
Xantusia vigilis Baird
Desert Night Lizard
OrIGINAL DescripTion.—Yantusia vigilis Baird, Proc. Acad. Nat.
Sei. Phila., 1858 [1859], p. 255.
Type Locaniry—Fort Tejon, California [probably Antelope
Valley, in northern Los Angeles County (see Grinnell and Grinnell,
Throop Inst. Bull., 35, 1907, p. 59) |.
Common Name.—Xantus’s Lizard.
Ranee.—Restricted to the tree yucea belt, chiefly on the Mohave
Desert. Ranges north to east slope of Inyo Mountains, Inyo County
(Meek, Field Columb. Mus., zool. ser., 7, 1905 [1906], p. 13) ; west in
vicinity of Walker Pass, Kern County, to Kelso Creek Valley, 3200
feet altitude, near Weldon (Mus. Vert. Zool.), to head of Piru Creek
(Meek, loc. cit.) and Pallett (Mus. Vert. Zool.), in northern Los
Angeles County, to Hesperia, San Bernardino County (Stejneger,
N. Amer. Fauna, 7, 1893, p. 198), and to Cabezon, Riverside County
(Van Denburgh, Proe. Calif. Acad. Sci., ser. 2, 5, 1895, p. 526) ; east
to Goffs [Blake P. O.], San Bernardino County (Camp, Univ. Calif.
Publ. Zool., 12, 1916, p. 528). Oceupies the Lower Sonoran life-zone.
Lives in and beneath prostrate trunks of tree yuceas.
Xantusia henshawi Stejneger
Henshaw Night Lizard
OrIGINAL DEscripTION.—Xantusia henshawi Stejneger, Proce. U.S.
Nat. Mus., 16, 1893 [1894], p. 467.
Tyrr Locauiry—Witch Creek, 2700 feet altitude, San Diego
County, California.
Synonyms.—NVantusia picta Cope, Amer. Nat., 29, 1895, pp. 859-
860, 939 (type from ‘*‘Tejon Pass’’—almost certainly Poway, San
Diego County: Van Denburgh, Copeia, no. 27, February 24, 1916,
pp. 14-15) ; Zablepsis henshawt.
Common Names.—Henshaw’s Lizard; Cope’s Lizard.
RancEe.—Known only from eastern San Diego County, at the type
locality as above, at Poway (Van Denburgh, loc. cit.), and in La
172 University of California Publications in Zoology [Vou.17
Puerta Valley (U.S. Nat. Mus.; Stephens, MS). Occurs in the Upper
Sonoran life-zone. Lives in crevices of rock outcrops.
Xantusia riversiana Cope
Island Night Lizard
OriainaL DescripTrion.—NXantusia riversiana Cope, Proe. Acad.
Nat. Sci. Phila., 1883 [1884], pp. 29-32 (see also Amer. Nat., 13, 1879,
p. 801).
Type Locauiry.—California; fixed as San Nicolas Island, Ventura
County, California, by Rivers (Amer. Nat., 23, 1889, p. 1100).
Common Name.—Rivers’s Lizard.
Rance.—Known only from San Nicolas Island, Santa Barbara
Island, and San Clemente Island. The record from Santa Catalina
Island (Van Denburgh, Proe. Calif. Acad. Sei., ser. 3, zool., 4, 1905,
p- 16) is now questioned (Van Denburgh, Proe. Calif. Acad. Sei.,
ser. 4, 4, 1914, p. 133). Lives beneath stones and in wind-drifted
debris at bases of bushes.
Family TEIIDAE
Cnemidophorus tigris tigris Baird and Girard
Desert Whip-tailed Lizard
OrIGINAL DescripTiION.—Cnenuidophorus tigris Baird and Girard,
Proe. Acad. Nat. Sei. Phila., 6, 1852, p. 69.
Type Locauiry.—Valley of the Great Salt Lake, Utah.
Synonyms.—Cnemidophorus gracilis Baird and Girard, Proce.
Acad. Nat. Sci. Phila., 6, 1852, p. 128 (type from desert of Colorado) ;
Cnemidophorus tessellatus tigris, part; Cnemidophorus tessellatus
tesscllatus, part; Cnemidophorus sexlineatus var. tesselatus ; Cnemi-
dophorus sexlineatus var. bocourtii Boulenger, Cat. Lizards Brit. Mus.,
2nd ed., 2, 1885, p. 367 (types from California).
Common Names.—Desert Whip-tail; Swift Jack; Tessellated
Lizard; Tiger Armor-bearer; Tiger Lizard.
Rance.—Deserts of southeastern California. Extends north to
Benton, Mono County (Mus. Vert. Zool.) ; west to Gray’s (altitude
6000 feet, near Kearsarge Pass) and Carroll Creek, in Inyo County
(Mus. Vert. Zool.), and to desert bases of San Gabriel, San Bernardino,
San Jacinto and other mountain ranges to the southward. Intergrades
with C. t. mundus through Walker Pass, and with C. t. stejnegeri in
1917] Grinnell-Camp: Amphibians and Reptiles of California 173
Antelope Valley, Los Angeles County, around the northeast base of
the San Jacinto Mountains, and east of the desert divides in western
Imperial County (Mus. Vert. Zool.). Oceupies the Lower Sonoran
life-zone. Inhabits sandy or gravelly ground both among rocks and
bushes and in open country. (See fig. 10.)
Cnemidophorus tigris tigris |
Intergrade
Cnemidophorus tigris mundus
Intergrade (tigris and stejnegeri)
Cnemidophorus tigris stejnegeri
Cnemidophorus hyperythrus beldingi |
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNLA.
Fig. 10. Distribution of Whip-tailed Lizards (Cnemidophorus) in California.
Cnemidophorus tigris mundus Camp
California Whip-tailed Lizard
OricInaL Derscription—Cnemidophorus tigris mundus Camp,
Uniy. Calif. Publ. Zool., 17, December 28, 1916, p. 71 (new name to
replace Cnemidophorus undulatus Hallowell, Proc. Acad. Nat. Sei.
Phila., 7, 1854, p. 94).
174 University of California Publications in Zoology [Vou.17
Type Locauiry.—Fort Yuma, in San Joachim Valley; later cor-
rected to Fort Miller, Fresno County, California (Stejneger, N. Amer.
Fauna, 7, 1893, p. 201).
Synonyms.—Cnemidophorus tessellatus tigris, part; Cnemidopho-
rus tessellatus tessellatus, part; Cnemidophorus tigris undulatus, part.
Common Names.—California Whip-tail; Tessellated Tiger Lizard.
RanGE.—Sacramento and San Joaquin valleys and adjacent moun-
tain slopes and detached valleys. Occurs north as far as MeCloud
River, in Shasta County (Cope, Ann. Rep. U. S. Nat. Mus., 1898
[1900], p. 579) ; west to Winslow, Glenn County (Mus. Vert. Zool.),
to Kelseyville, Lake County (Van Denburgh, Oce. Papers Calif. Acad.
Sei., 5, 1897, p. 188), to Mount Diablo, Contra Costa County (Mus.
Vert. Zool.), to Los Gatos, Santa Clara County (Van Denburgh,
loc. cit.), to Carmel Valley, Monterey County (MeLain, Crit. Notes
Coll. Reptiles Western U. S., 1899, p. 9), and to Santa Margarita,
San Luis Obispo County (Mus. Vert. Zool.) ; south to Walker, Te-
hachapi, and Tejon passes, in Kern and Los Angeles counties; east to
Coulterville road near Big Meadow, 4000 feet altitude, in Mariposa
County (Storer, MS), and to Raymond, Madera County (Mus. Vert.
Zool.). Probably intergrades on the south with C. ¢t. stejnegeri.
Oceupies the Lower and Upper Sonoran life-zones. Usual habitat,
open valley floors; also lives amid sparse chaparral on hillsides. (See
fig. 10.)
Cnemidophorus tigris stejnegeri Van Denburgh
Stejneger Whip-tailed Lizard
OrIGINAL DEscRIPTION.—Cnemidophorus stejnegert Van Denburgh,
Proce. Calif. Acad. Sci., ser. 2, 4, July 12, 1894, pp. 300-301.
Type Locariry.—Between San Rafael and Ensenada, Lower Cali-
fornia, Mexico.
Synonyms.—Cnemidophorus grahamii stejnegerti; Cnemidophorus
tessellatus tessellatus, part; Cnemidophorus grahamit; Cnenudophorus
tigris undulatus, part.
Common Names.—Graham’s Striped Lizard; Stejneger’s Whip-
tail.
RANGE.
Chiefly Pacifie slope of southern California. Occurs
north to Matilija, Ventura County (Mus. Vert. Zool.) ; east to San
Gabriel Mountains, up to 5900 feet altitude, in Los Angeles County
(Camp, MS), to Lytle Creek (Van Denburgh, Oce. Papers Calif.
1917] Grinnell-Camp: Amphibians and Reptiles of California 175
Aead. Sci., 5, 1897, p. 140) and Upper Santa Ana Cafion, 5500 feet
altitude (Grinnell, Univ. Calf. Publ. Zool., 5, 1908, p. 163), in San
Bernardino County, to San Jacinto and Santa Rosa mountains, River-
side County (Atsatt, Univ. Calif. Publ. Zool., 12, 1913, pp. 39-40),
and to Vallecito, in eastern San Diego County (Mus. Vert. Zool.).
Occupies the Lower and Upper Sonoran life-zones. Inhabits washes,
gravelly mesas, and dry mountain slopes. (See fig. 10.)
Cnemidophorus hyperythrus beldingi (Stejneger)
Belding Orange-throated Lizard
Oriemwnau Description.—Verticaria beldingi Stejneger, Proc. U.S.
Nat. Mus., 17, 1894 [1895], pp. 17-18.
Type Locautrry.—Cerros Island, Lower California, Mexico.
Synonyms.—Verticaria hyperythra; Cnemidophorus hyperythrus ;
Verticaria hyperythra beldingt.
Common Names.—Belding’s Orange-throat ; Cape Striped Lizard.
Rance.—Foothill districts of western San Diego and Riverside
counties. Occurs north to Reche Canon, Riverside County (Atsatt,
Uniy. Calif. Publ. Zool., 12, 1913, p. 40) ; west to Temescal Mountains,
Riverside County (Van Denburgh, Proe. Calif. Acad. Sei., ser. 4, 3,
1912, p. 150), and to Escondido (Mus. Vert. Zool.) and San Diego
(Cope, Ann. Rep. U. 8. Nat. Mus., 1898 [1900], p. 564), in San Diego
County; east to Oak Grove, San Diego County (Van Denburgh, loc.
cit., p. 152), and San Jacinto, Riverside County (Stejneger, loc. cit.).
Occurs within the Lower Sonoran life-zone. Inhabits dry, sparsely
vegetated, sandy ground, and dusty roadsides. (See fig. 10.)
Family SCINCIDAE
Plestiodon skiltonianum Baird and Girard
Western Skink
OriGINAL DeEscription.—Plestiodon Skiltonianuwm Baird and
Girard, Proe. Acad. Nat. Sei. Phila., 6, 1852, p. 69.
Type Locauiry.—Oregon.
Synonyms.—Eumeces quadrilineatu[s| Hallowell, Pac. R. R. Rep.,
10, 1859, Williamson’s Rep., pt. 4, no. 1, p. 10 (types from near
Mohave River and in San Bernardino Valley [more lkely the latter] ) ;
Eumeces gilberti Van Denburgh, Proe. Calif. Acad. Sei., ser. 2, 6, 1896,
176 University of California Publications in Zoology [Vou.17
pp. 350-352 (type from Yosemite Valley, Mariposa County [see Camp,
Univ. Calif. Publ. Zool., 17, 1916, pp. 72-73] ) ; Hwmeces skiltonvanus
var. brevipes Cope, Ann. Rep. U. S. Nat. Mus., 1898 [1900], pp. 648-
644 (type from Fresno) ; Hwmeces hallowellii Bocourt, in Duméril,
Bocourt, and Moquard, in Miss. Sei. au Mex., Recherch. Zool., pt. 3,
sec. 1, 6° livr., 1879, p. 485, Atlas, [1881], pl. 22n, fig. 7 (type from
California) ; Humeces skiltonianus var. amblygrammus Cope, loc. cit.,
p. 648 (type from Fort Humboldt) ; Humeces skiltonianus.
Common Names.—Blue-tailed Lizard; Skilton’s Skink; Red-headed
Skink; Gilbert’s Skink; Blue-tailed Skink.
RanGE.—Nearly the entire state; absent on the southeastern deserts
and on the Sierra Nevada above 8000 feet altitude. Occurs east to
head of Willow Creek, 7000 feet altitude, Panamint Mountains, Inyo
County (Stejneger, N. Amer. Fauna, 7, 1893, p. 202) ; southeast to
Maturango Spring, Argus Range, Inyo County (Stejneger, loc. cit.) ;
to Kern River near Isabella (Mus. Vert. Zool.) and old Fort Tejon
(Stejneger, loc. cit.), in Kern County; to Barley Flats, 5500 feet
altitude, San Gabriel Mountains, Los Angeles County (Mus. Vert.
Zool.) ; to Bluff Lake, San Bernardino Mountains, San Bernardino
County (Van Denburgh, Proce. Calif. Acad. Sci., ser. 4, 3, 1912,
p. 149) ; to Strawberry Valley, 5500 feet altitude, San Jacinto Moun-
tains, Riverside County (Atsatt, Univ. Calif. Publ. Zool., 12, 1913,
p. 41) ; and to Jacumba Hot Springs, San Diego County (Cope, Ann.
Rep. U. S. Nat. Mus., 1898 [1900], p. 644). Occurs also on Santa
Catalina Island (Van Denburgh, Proce. Calif. Acad. Sci., ser. 4, 4, 1914,
pp. 132, 138). Inhabits chiefly the Upper Sonoran life-zone, but
extends also through Transition. Characteristic habitat, rocky or
wooded hillsides and cafon bottoms; lives beneath stones and masses
of dead vegetation.
Suborder SERPENTES
Family LEPTOTYPHLOPIDAE
Leptotyphlops humilis (Baird and Girard)
Worm Snake
ORIGINAL DescripTION.—Rena humilis Baird and Girard, Cat. N.
A. Reptiles in Smiths. Inst., pt. 1, 1853, p. 143.
Type Locaurry.—Valliecitas [— Vallecito, San Diego County],
California.
~]
1917] Grinnell-Camp: Amphibians and Reptiles of California 1
Synonyms.—[?] Ophisaurus ventralis; Glauconia humilis; Steno-
stoma humile; Siagonodon humilis.
Common Names.—California Rena; California Blind Snake; Glass
Snake ; Sheep-nosed Snake.
Ranee.—The three definite records are from southeastern Cali-
fornia: six miles from Bennett Wells, in Death Valley, Inyo County
(Stejneger, N. Amer. Fauna, 7, 1893, p. 203); San Bernardino, San
Bernardino County (Boulenger, Cat. Snakes Brit. Mus., 3, 1896,
p. 591); and the type locality, in extreme eastern San Diego County,
as above. These localities lie within the Lower Sonoran life-zone.
Family BOIDAE
Subfamily Bomar
Lichanura roseofusca Cope
California Boa
ORIGINAL Descriprion.—Lichanura roseofusca Cope, Proce. Acad.
Nat. Sci. Phila., 1868, p. 2.
Type Locauiry.—Northern Lower California, Mexico.
Synonyms.—Lichanura orcutti Stejneger, Proce. U. S. Nat. Mus.,
12, 1889, pp. 96-97, fig. 1 (type from Colorado Desert, San Diego
County); Lichanura simplex Stejneger, loc. cit., pp. 97-99, fig. 2
(type from San Diego) ; Lichanura myriolepis; Lichanura trivirgata.
Common Names.—Rubber Snake, part; Rubber Boa, part; Rosy
Boa.
Rance.—Limited to a small area in the southern part of the state.
Recorded northwest to Arroyo Seco (Camp, MS) and Mount Wilson
(Van Denburgh, Oce. Papers Calif. Acad. Sci., 5, 1897, p. 154) near
Pasadena, Los Angeles County; east to Cabezon and Palm Canon,
Riverside County (Atsatt, Univ. Calf. Publ. Zool., 12, 1913, p. 41),
and to Dulzura (Mus. Vert. Zool.) and ‘‘ Colorado Desert’’ (as above),
San Diego County. There is a specimen in the U.S. National Museum
(no. 44317) from the Providence Mountains, northeastern San Ber-
nardino County. Occurs within the Lower and Upper Sonoran life-
zones. Lives on shaded hillsides in dense chaparral. (See fig. 11.)
Charina bottae (Blainville)
Rubber Snake
OrteInAL Description.—Tortrix Bottae Blainville, Nouv. Ann.
Mus. Hist. Nat., 4, 1835, pp. [57-58] 289-290, pl. 26, figs. 1, la, 1b.
178 University of California Publications in Zoology [Vou.17
Type Locauiry.—California [Monterey ?].
Synonyms.—Charina brachyops Cope, Proc. U. S. Nat. Mus., 11,
1888, p. 88 (type from Point Reyes, Marin County) ; Charina plumbea;
Pseudoeryx bottae.
=
eae
Ie SS a “| | y
Ate! @ i }
ty
oy : Lichanura roseofusca
Charina bottae
Rhinocheilus lecontei
Salvadora hexalepis
+b bde@O
Hypsiglena ochrorhynchus
Ranges of Charina bottae and Lichanura
roseofusca outlined
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA cS
Fig. 11. Distribution of Snakes (Lichanura, Charina, Rhinocheilus, Salvadora,
and Hypsiglena) in California.
Common Names.—Two-headed Snake; Lead-colored Worm Snake ;
Wood Snake; Rubber Boa, part.
Ranee.—Northern California, both east and west of the Sierra
Nevada. Has been found south to Redwood Cafion, East Fork Kaweah
River, Tulare County (Stejneger, N. Amer. Fauna, 7, 1893, p. 203),
1917] Grinnell-Camp: Amphibians and Reptiles of California 179
and to Soquel, Santa Cruz County (Van Denburgh, Occ. Papers Calif.
Acad. Sci., 5, 1897, p. 156). Southeasternmost stations are: Tahoe
City, Placer County (Van Denburgh, loc. cit.), and Yosemite Valley,
Mariposa County (Mus. Vert. Zool.). Occurs chiefly within the
Transition life-zone, extending locally into Upper Sonoran. Inhabits
damp ground in canons and on forest floors. (See fig. 11.)
Superfamily COLUBROIDEAE
Family COLUBRIDAE
Subfamily NarricinaE
Thamnophis marcianus (Baird and Girard)
Marey Garter Snake
OriginaL Description.—Eutainia Marciana Baird and Girard,
Cat. N. A. Reptiles in Smiths. Inst., pt. 1, 1853, pp. 36-37.
Type Locaurry—Red River, Arkansas [=-near Cache Creek,
Oklahoma: Ruthven, U. S. Nat. Mus. Bull., 61, 1908, p. 58].
SynonyM.—Lutaenia elegans marciana.
Rance—Along the lower Colorado River from Fort Yuma, In-
perial County (Yarrow, U. S. Nat. Mus. Bull, 24, 1882, pp. 17, 118)
north at least to Riverside Mountain, Riverside County (Mus. Vert.
Zool.). Life-zone, Lower Sonoran. Habitat, riparian.
Thamnophis ordinoides ordinoides (Baird and Girard)
Pacifie Coast Garter Snake
ORIGINAL Description —Tropidonotus ordinoides Baird and Gi-
rard, Proc. Acad. Nat. Sci. Phila., 6, 1852, p. 176.
Type Locauiry.—Puget Sound.
Synonyms.—Eutaenia leptocephala; Eutaenia infernalis vidua
Cope, Proc. U. S. Nat. Mus., 14, 1891, pp. 658-659 (types from San
Francisco) ; Eutainia atrata Kennicott, in Cooper, Pace. R. R. Rep.,
12, 1859, pt. 3, no. 4, p. 296 (type from California) ; Hutaenia elegans,
part; Eutaenia infernalis, part; Eutaenia elegans ordinoides; Tropi-
donotus leptocephalus; Eutaenia sirtalis leptocephala; Eutaenia ordi-
noides; Thamnophis infernalis, part; Eutainia sirtalis elegans;
Thamnophis leptocephala.
Common Names.—Puget Garter Snake; Narrow-headed Garter
Snake; Black Garter Snake; Boyd’s Garter Snake, part.
Rance.—Chiefly the narrow northwest coast strip within twenty
180 University of California Publications in Zoology [Vou.17
miles of the seacoast ; authentically recorded south as far as Monterey
(Ruthven, U. 8. Nat. Mus. Bull., 61, 1908, p. 149). One interior
record: Fresno (Ruthven, loc. cit.). Occupies the Transition life-zone.
Lives in dense vegetation, along streams and on marshy ground.
Thamnophis ordinoides elegans (Baird and Girard)
Elegant Garter Snake
OriginaL DescripTion.—Eutainia elegans Baird and Girard, Cat.
N. A. Reptiles in Smiths. Inst., pt. 1, 1853, pp. 84-35.
Typr Locaurry.—Eldorado County, California.
Synonyms.—Hutaina vagrans Baird and Girard, loc. cit., pp. 35-
36 (types from Humboldt River) ; Lutaenia infernalis, part ; Eutaena
elegans lincolata Cope, Proe. U. S. Nat. Mus., 14, 1891, p. 655 (type
from southern California: see Cope, Ann. Rep. U. S. Nat. Mus., 1898
[1900], p. 1039) ; Hutaenia elegans brunnea Cope, Proce. U. S. Nat.
Mus., 14, 1891, p. 654 (type from Fort Bidwell, Modoe County) ;
Eutacnia couchti Kennicott, in Baird, Pac. R. R. Rep., 10, 1859, Ab-
bot’s Rep., pt. 4, no. 4, pp. 10-11 (type from bank of Pit River) ;
Thamnophis infernalis, part; Tropidonotus tri-vittatus Hallowell,
Proce. Acad. Nat. Sci. Phila., 6, 1853, p. 237 (types from banks of
Cosumnes and other rivers in California) ; Eutaenia elegans couchi,
part; Tropidonotus ordinatus infernalis; Tropidonotus ordinatus vay.
couchti; Thamnophis vagrans; Thamnophis parietalis, part; Eutaenia
elegans vagrans; Eutaenia hammondi, part; Thamnophis elegans,
part; Hutaenia elegans infernalis.
Common Names.—Boyd’s Garter Snake, part; Pacifie Garter
Snake, part; Wandering Garter Snake; Hammond’s Garter Snake,
part; Single-striped Garter Snake; Green Garter Snake; Western
Garter Snake, part.
Rance.—Northeastern portion of the state; across the head of the
Sacramento Valley and reaching in that latitude as far as Humboldt
Bay ; south along the Sierra Nevada to Lone Pine, Inyo County, and
Kern River, in Kern County, and thence west through the mountains
about the head of the San Joaquin Valley to the seacoast from Santa
Ynez River, Santa Barbara County (Mus. Vert. Zool.), north to Morro,
San Luis Obispo County (Ruthven, U. S. Nat. Mus. Bull., 61, 1908,
pp. 140-141). Thought to intergrade along the western edge of its
range with ordinoides and at the south with hammondu. Occurs
within the Canadian and Transition life-zones, extending locally into
Upper Sonoran. Inhabits marshes, stream-sides, and ponds.
1917] Grinnell-Camp: Amphibians and Reptiles of California 181
Thamnophis ordinoides hammondii (Kennicott)
California Garter Snake
Ori@inaL Description.—Eutaenia Hammondii Kennicott, Proc.
Acad. Nat. Sei. Phila., 1860 [1861], p. 332.
Tyre Locauiry.—San Diego, California.
Synonyms.—LHutaenia couchu, part; Eutaenia elegans couch,
part; Tropidonotus ordinatus var. hammondu; Tropidonotus ordi-
natus; Eutaenia marciana, part; Tropidonotus vagrans.
Common Names.—Hammond’s Garter Snake, part; Water Snake;
Couch’s Garter Snake; Marey’s Garter Snake, part.
RanGE.—The southwestern portion of the state, chiefly in moun-
tains west of the desert divides. Recorded north to Lone Pine, Inyo
County (U. S. Nat. Mus.), to Kernville, Kern County, and vicinity
of Fresno (Ruthven, U. 8. Nat. Mus. Bull., 61, 1908, p. 135). East-
ernmost stations are: Laguna Mountains, San Diego County (Cope,
Ann. Rep. U. 8. Nat. Mus., 1898 [1900], p. 1044) ; Palm Canon, 800
feet altitude, San Jacinto Mountains (Atsatt, Univ. Calif. Publ. Zool.,
12, 1913, p. 43); Bluff Lake, 7500 feet altitude, San Bernardino
Mountains (Mus. Vert. Zool.) ; and Mohave River at Victorville (Mus.
Vert. Zool.). Oceupies the Upper Sonoran and Transition life-zones.
Inhabits streams and margins of lakes and the near vicinity of same.
Thamnophis sirtalis parietalis (Say)
Western Garter Snake
ORIGINAL DeEscripTion.—Coluber parietalis Say, in Long, Exped.
to Rocky Mountains, 1, 1823, pp. 186-187, footnote.
Type Locaurry.—Camp Missouri [near Council Bluffs, Iowa].
Synonyms.—Coluber infernalis Blainville, Nouv. Ann. Mus. Nat.
Hist., 4, 1835, pp. [59-60] 291-292, pl. 26, figs. 3-3a (type from Cali-
fornia); Eutaenia sirtalis obscura; Eutaenia sirtalis; Tropidonotus
parietalis; Eutaenia proxima; Eutaenia imperialis (type said to be
from Tomales Bay: Coues and Yarrow, Bull. U. 8. Geol. Surv. Terr.,
4, 1878, p. 280) ; Eutaenia sirtalis pickeringti; Thamnophis infernalis,
part; Thamnophis elegans, part; Eutaenia concinna; Thamnophis
parietalis; Eutaenia sirtalis parietalis; Eutaena sirtalis tetrataena;
Eutaenia sirtalis dorsalis.
Common Names.—Pacific Garter Snake, part; Rocky Mountain
Garter Snake; Red-barred Garter Snake; California Garter Snake,
182 University of California Publications in Zoology [Vou.17
part; Churchill’s Garter Snake; Dusky Garter Snake; Say’s Garter
Snake; Striped Snake; Pickering’s Garter Snake.
Ranee.—Almost throughout the state west and north of the south-
eastern deserts. Recorded southeast to Yosemite Valley, Mariposa
County (Van Denburgh, Occ. Papers Calif. Acad. Sei., 5, 1897, p. 203),
to Kern River, in Kern County (Mus. Vert. Zool.), and to Riverside,
Riverside County (Van Denburgh, loc. cit.). Southernmost station,
Bixby, near Long Beach, Los Angeles County (Grinnell and Grinnell,
Throop Inst. Bull, 35, 1907, pp. 48-49). Oceupies the Lower and
Upper Sonoran and Transition life-zones. Lives along streams,
sloughs, and lake-margins, and in wet meadows.
Subfamily CoLuBRINAE
Chilomeniscus cinctus Cope
Banded Burrowing Snake
OrieInAL Description.—Chilomeniscus cinctus Cope, Proe. Acad.
Nat. Sei. Phila., 1861, p. 303.
Type Locaniry.—Near Guaymas, east coast of Gulf of California,
Mexico.
Synonym.—Chilomeniscus ephippicus Cope, Proe. Acad. Nat. Sci.
Phila., 1867, p. 85 (type from Owens Valley, Inyo County).
Common Names.—Horse Snake; Red and Black Ground Snake.
Rance.—Only two records, both from the southeastern deserts:
from the type locality of ephippicus, as above, and from Fort Yuma,
Imperial County (Van Denburgh and Slevin, Proce. Calif. Acad. Sei.,
ser. 4, 3, 1913, p. 410). Life-zone, Lower Sonoran. (See fig. 12.)
Sonora occipitalis (Hallowell)
Desert Burrowing Snake
OrigINAL DescripTion.—Rhinostoma occipitale Hallowell, Proe.
Acad. Nat. Sei. Phila., 7, 1854, p. 95.
Type Locatiry.—Mohave Desert, California.
Synonyms.—Lamprosoma annulatum Baird, U. S. Mex. Bound.
Surv., 2, 1859, pt. 2, Reptiles, p. 22, pl. 21, fig. 1 (types from Colorado
Desert) ; Lamprosoma occipitale; Chionactis occipitalis; Chionactis
occipitalis annulatus.
Common Names.—Desert Snake; Mohave Ringed Snake.
1917] Grinnell-Camp: Amphibians and Reptiles of California 183
RanGe.—Mohave and Colorado deserts, north to Owens Lake, Inyo
County (Meek, Field Columb. Mus., zool. ser., 7, 1906, p. 15), and
Blythe Junction, San Bernardino County (Camp, Univ. Calif. Publ.
Zool., 12, 1916, p. 531), and west to Carrizo Creek and La Puerta
Valley, in eastern San Diego County (Mus. Vert. Zool.). Restricted
to the Lower Sonoran life-zone. Inhabits open gravelly or sandy
ground. (See fig. 12.)
Sonora episcopa (Kennicott)
Texas Ground Snake
OrigInaL Derscription.—Lamprosoma episcopum Kennicott, in
Baird, U. S. Mex. Bound. Surv., 2, 1859; pt. 2, Reptiles, p. 22, pl. 8,
fig. 2.
Type Locauiry.—Eagle Pass, Texas.
RanGe.—Extreme southeastern deserts; two instances of occur-
rence: 4 miles north of Blythe Junction, San Bernardino County
(Camp, Univ. Calif. Publ. Zool., 12, 1916, pp. 530-531) ; Heber, Im-
perial County (no. 5610, Mus. Vert. Zool.). Both localities he within
the Lower Sonoran life-zone. In the first case, the snake was taken
from beneath a stone on a rocky hillside. (See fig. 12.)
Contia mitis Baird and Girard
Sharp-tailed Snake
OriGINAL DEscrIeTION.—Contia mitis Baird and Girard, Cat. N.
A. Reptiles in Smiths. Inst., pt. 1, 1853, pp. 110-111.
Tyre Locaniry.—San Jose, Santa Clara County, California.
Synonyms.—Ablabes purpureocauda Giinther, Cat. Colubrine
Snakes Brit. Mus., 1858, p. 245 (type from California) ; Homalosoma
mite.
Common Names.—Purple-tailed Snake; Pacifie Ground Snake;
Brown Snake; Gentle Brown Snake.
RaNnce.—Northern portion of the state, mostly near the seacoast.
Southernmost stations: Big Basin, Santa Cruz County (Van Den-
burgh, Oce. Papers Calif. Acad. Sei., 5, 1897, p. 163), and [near]
Fresno, Fresno County (Yarrow, U. S. Nat. Mus. Bull., 24, 1882,
pp. 14, 87) ; easternmost stations: Fyffe, Eldorado County (Van Den-
burgh, loc. cit.), and Baird, Shasta County (Cope, Ann. Rep. U. 8.
Nat. Mus., 1898 [1900], p. 926). Occurs in the Transition and Upper
Sonoran life-zones. Lives beneath stones in hilly country. (See
fig. 12.)
184 University of California Publications in Zoology [Vou.17
Diadophis amabilis Baird and Girard
Western Ring-necked Snake
ORIGINAL DeEscripTioN.—Diadophis amabilis Baird and Girard,
Cat. N. A. Reptiles in Smiths. Inst., pt. 1, 1853, pp. 113-114.
Type Locauiry—San Jose, Santa Clara County, California.
Synonyms.—Diadophis pulchellus Baird and Girard, loc. cit., p.
115 (type from Eldorado County) ; Diadophis punctatus pulchellus ;
Diadophis punctatus amabilis; Diadophis amabilis pulchellus ; Coro-
nella amabilis; Ablabes punctatus; [2] Coluber punctatus; Diadophis
punctatus.
Common Names.—California Ring-necked Snake; Red-bellied
Snake; Spotted Ring Snake.
Rance—West of the desert divides the whole length of the state.
Occurs northeast to McCloud River, Shasta County (Townsend, Proe.
U.S. Nat. Mus., 10, 1887, p. 239) ; east to Oroville, Butte County (Mus.
Vert. Zool.) ; to Eldorado County (as above); to Yosemite Valley,
Mariposa County (Stejneger, N. Amer. Fauna, 7, 1893, p. 204) ; to
[near] Fresno, Fresno County (Yarrow, U. 8. Nat. Mus. Bull, 24,
1882, pp. 15, 95-96); to Tejon Pass [Kern County?] (Heermann,
Pac. R. R. Rep., 10, 1859, Williamson’s Rep., pt. 4, no. 1, p. 24) ; to
Arroyo Seco Cation (Grinnell and Grinnell, Throop Inst. Bull., 35,
1907, pp. 38-39) and Glendora (Mus. Vert. Zool.), Los Angeles
County; to Santa Ana Cafion, 6400 feet altitude, San Bernardino
County (Grinnell, Univ. Calif. Publ. Zool., 5, 1908, p. 164) ; to Straw-
berry Valley, 5500 feet altitude, San Jacinto Mountains, Riverside
County (Atsatt, Univ. Calif. Publ. Zool., 12, 1913, p. 41); and to
Witch Creek, San Diego County (Cope, Ann. Rep. U. 8. Nat. Mus.,
1898 [1900], p. 749). There is a mutilated specimen of a Diadophis
in the U. S. National Museum from Santa Catalina Island. Occurs
within the Upper Sonoran and Transition life-zones. Inhabits shaded
canons; lives in masses of dead leaves and beneath stones. (See
fio 25)
Lampropeltis pyromelana multicincta (Yarrow)
Coral King Snake
OrIGINAL Description.—Ophibolus getulus multicinctus Yarrow,
Proce. U. S. Nat. Mus., 5, 1882, p. 440.
Type Locauiry.—|Near] Fresno, California.
Synonyms.—[?] Coluber (Zacholus) zonatus Blainville, Nouv.
1917] Grinnell-Camp: Amphibians and Reptiles of California 185
Ann. Mus. Nat. Hist., 4, 1835, pp. [61-62] 293-294 (type from Cali-
fornia) ; Coronella multifasciata Bocourt, in Duméril, Bocourt, and
Moquard, in Miss. Sci. au Mex., Recherch. Zool., pt. 3, see. 1, 10°, livr.
1886, pp. 616-617, Atlas, pl. 40, figs. 2-2c (type from California) ;
eRe ee =
Chilomeniscus cinctus
Sonora occipitalis
Sonora episcopa
Contia mitis
Diadophis amabilis
Tantilla eiseni
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
“ALLPORNLA
UNIVERSITY OF
Fig. 12. Distribution of Snakes (Diadophis, Tantilla, Chilomeniscus, Sonora,
and Contia) in California.
Bellophis zonatus Lockington, Proc. Calif. Acad. Sci., 7, 1876 [1877],
pp. 52-53 (type from ‘‘Northern California’’: Santa Barbara, ac-
cording to Van Denburgh, Occ. Papers Calif. Acad. Sei., 5, 1897,
p. 167) ; Ophibolus pyrrhomelanus ; Ophibolus pyrrhomelas; Coronella
pyromelanus zonata; Coronella zonata; Ophibolus zonatus; Lampro-
peltis zonata.
186 University of California Publications in Zoology (Vou. 17
Common Names.—California King Snake; Arizona Ringed Snake ;
Ringed King Snake; Hisen’s King Snake; Red Milk Snake; Corral
Snake; Ring Snake; Harlequin Snake.
RanGe.—The southwestern portion of the state, altogether west
of the desert divides. Occurs north in the coast belt as far as Glen-
wood, Santa Cruz County, and Mount Hamilton, Santa Clara County
(Van Denburgh, loc. cit., p. 169), and on the west slope of the Sierra
Nevada to Riverton, Eldorado County (Van Denburgh, loc. cit.) ; east
to Yosemite Valley, Mariposa County (Van Denburgh, loc. cit.) ; to
Heaven’s Gate, near Little Kern Lake, Tulare County (Van Den-
burgh, loc. cit.) ; to Arroyo Seco Canon, near Pasadena, Los Angeles
County (Mus. Vert. Zool.) ; to upper Santa Ana Cafion, 5500 feet
altitude, San Bernardino County (Grinnell, Univ. Calif. Publ. Zool.,
5, 1908, p. 165) ; and to Strawberry Valley, 6000 feet altitude, River-
side County (Atsatt, Univ. Calif. Publ. Zool., 12, 1913, pp. 41-42).
Extends south as far as vicinity of San Diego (Van Denburgh, loc.
cit.). Oceurs chiefly within the Transition life-zone, entering to some
extent the Upper Sonoran. Inhabits forest floors and chaparral-
covered hillsides.
Lampropeltis boylii (Baird and Girard)
Boyle King Snake
OrtGInaAL Description.—Ophibolus Boylit Baird and Girard, Cat.
N. A. Reptiles in Smiths. Inst., pt. 1, 1853, pp. 82-83.
Type Locaniry.—Eldorado County, California.
Synonyms.—Coronella balteata Hallowell, Proce. Acad. Nat. Sei.
Phila., 6, 1853, pp. 236-237 (types from California) ; Ophibolus
getulus boyli, part; Coronella getula, part; Coronella boylit.
Common NameEs.—Boyle Milk Snake ; California King Snake, part;
California Milk Snake, part; Banded Milk Snake.
Rance.—Throughout the southern and central parts of the state,
except on the high mountains (above 6000 feet altitude) and along
the lower Colorado River. Occurs north in the coast belt at least to
near Cazadero, Sonoma County (Mus. Vert. Zool.), in the interior to
Lierly’s, near Mount Sanhedrin, Mendocino County (Mus. Vert. Zool.),
to McCloud River, Shasta County (Townsend, Proc. U. S. Nat. Mus.,
10, 1887, p. 239), and to Applegate, Placer County (Van Denburgh,
Oce. Papers Calif. Acad. Sei., 5, 1897, p. 171), and, east of the Sierran
divide, to Beveridge Canon (on east slope of Inyo Mountains) and
1917] Grinnell-Camp: Amphibians and Reptiles of California 187
Wild Rose Springs, in Inyo County (Meek, Field Columb. Mus., zool.
ser., 7, 1905 [1906], p. 15). There is a specimen in the U. S. National
Museum from Avalon, Santa Catalina Island [see also Cooper, Proe.
Calif. Acad. Sci., 4, 1870, p. 79]. Occurs in the Lower and Upper
Sonoran and Transition life-zones. Shows no special restriction in
habitat.
Lampropeltis conjuncta Cope
Black King Snake
OriagwwaL Description.—Lampropeltis boylii var. conjuncta Cope,
Proc. Acad. Nat. Sei. Phila., 1861, pp. 301-302.
Type Locauiry.—Cape San Lueas [Lower California, Mexico].
SYNONYM.
Ophibolus getulus boyliv, part.
Common Name.—California King Snake, part.
Rance.—Colorado River bottom. Has been taken near Pilot Knob
and 5 miles northeast of Fort Yuma (nos. 1837 and 1838, Mus. Vert.
Zool.), and at Fort Yuma (Cope, loc. cit.). These stations are all in
Imperial County and le within the Lower Sonoran life-zone.
Lampropeltis californiae (Blainville)
California King Snake
OriGINAL DeEscription.—Coluber (Ophis) Californiae Blainville,
Nouv. Ann. Mus. Nat. Hist., 4, 1835, p. [60] 292, pl. 27, figs. 1, 1a, 1b.
Type Locauiry.—California.
Synonyms.—Ophibolus getulus eisent Yarrow, Proe. U. 8. Nat.
Mus., 5, 1882, pp. 489-440 (types from Fresno, California) ; Ophibolus
getulus californiae; Ophibolus californiae; Coronella Californiae ;
Coronella getula, part; Coronella getulus var. californica.
Common Names.—Blainville’s King Snake; California Milk Snake,
part.
Rance.—Interruptedly distributed through the southern part of
the state west of the desert divides. Localities of occurrence are:
Fresno (as above) ; Waterman Canon, San Bernardino County (Van
Denburgh, Proc. Calif. Acad. Sci., ser. 4, 3, 1912, p. 149) ; Riverside
County (Van Denburgh, Oce. Papers Calif. Acad. Sei., 5, 1897, p.
174) ; Cuyamaca (Van Denburgh, 1912, loc. cit., p. 151), Witch Creek
(Cope, Ann. Rep. U. 8. Nat. Mus., 1898 [1900], p. 923), Dulzura and
Julian (Mus. Vert. Zool.), in San Diego County. Occurs in the
Upper, and possibly also the Lower, Sonoran life-zone.
188 University of California Publications in Zoology (Vou. 17
Rhinocheilus lecontei Baird and Girard
Long-nosed Snake
OrIGINAL DEscRIPTION.—Rhinocheilus Lecontei Baird and Girard,
Cat. N. A. Reptiles in Smiths. Inst., pt. 1, 1853, pp. 120-121.
Type Locauiry.—San Diego, California.
Common Name.—Leconte’s Snake.
Ranee—Chiefly Pacific slope of southern California and floor of
San Joaquin Valley. Has been found northwest to Carrizo Plain,
San Luis Obispo County (Mus. Vert. Zool.), and to Fresno (Yarrow,
U.S. Nat. Mus. Bull., 24, 1882, pp. 14, 18). Hasternmost stations
are: Independence, Inyo County (Mus. Vert. Zool.), Pasadena, Los
Angeles County (Mus. Vert. Zool.), and Cabezon and Dos Palmos
Spring, 3500 feet altitude, Santa Rosa Mountains, in Riverside County
(Atsatt, Univ. Calif. Publ. Zool., 12, 1913, p. 42). Occurs within the
Lower Sonoran life-zone. Inhabits open flat country, living in rodent
burrows. (See fig. 11.)
Hypsiglena ochrorhynchus Cope
Spotted Night Snake
OriciInAL Derscription.—Hypsiglena ochrorhynchus Cope, Proce.
Acad. Nat. Sei. Phila., 1860, pp. 246-247.
Type Locaniry.—Cape San Lueas, [Lower] California.
Common Names.—Rock Snake, Xantus’s Snake.
Rance.—The southern portion of the state, chiefly in mountainous
districts. The stations of occurrence known to us are as follows:
Near Christy, Contra Costa County (Mus. Vert. Zool.) ; foothills near
Los Gatos, Santa Clara County (Van Denburgh, Proe. Calif. Acad.
Sci., ser. 3, zool., 4, 1906, pp. 65-66) ; Shepherd Canon, Argus Range,
Inyo County (Stejneger, N. Amer. Fauna, 7, 1893, p. 204) ; near Los
Angeles (Riithling, Copeia, no. 15, February 20, 1915); Hesperia
(Van Denburgh, Oce. Papers Calif. Acad. Sci., 5, 1897, p. 180) and
Santa Ana Canon, 5500 feet altitude (Grinnell, Univ. Calf. Publ.
Zool., 5, 1908, p. 165), in San Bernardino County ; Strawberry Valley,
5000 feet altitude, and San Jacinto, in Riverside County (Van Den-
burgh, loc. cit.) ; San Diego (Van Denburgh, loc. cit.), Witch Creek
(Cope, Ann. Rep. U. S. Nat. Mus., 1898 [1900], p. 954), and Cuyamaca
Mountains (Van Denburgh, loc. cit.), im San Diego County. Occurs
within the Lower and Upper Sonoran life-zones. Inhabits rocky
situations. (See fig. 11.)
1917] Grinnell-Camp: Amphibians and Reptiles of California 189
Salvadora hexalepis (Cope)
Patch-nosed Snake
OriegInAL Description.—Phimothyra hexalepis Cope, Proe. Acad.
Nat. Sci. Phila., 1866 [1867], p. 304.
Type Locaniry.—Fort Whipple, Arizona.
Synonyms.—Salvadora grahamiae hexalepis; Zamenis grahami;
Salvadora grahamiac.
Common Names.—Banded Flat-nosed Snake; Graham’s Flat-nosed
Snake.
Ranee.—The southern portion of the state, chiefly in arid situ-
ations. Has been taken north to Amargosa Borax Works and Matu-
rango Spring, Argus Range, in Inyo County (Stejneger, N. Amer.
Fauna, 7, 1893, p. 206); west to Arroyo Seco, near Pasadena, Los
Angeles County (Grinnell and Grinnell, Throop Inst. Bull., 35, 1907,
p. 42), to Riverside (McLain, Crit. Notes Coll. Reptiles Western U. S.,
1899, p. 11) and to San Diego (Cope, Ann. Rep. U. 8. Nat. Mus., 1898
[1900], p. 820). Oceupies the Lower and Upper Sonoran life-zones.
Inhabits hillsides with sparse covering of bushes. (See fig. 11.)
Coluber constrictor vetustus (Baird and Girard)
Western Yellow-bellied Racer
ORIGINAL Description.—Bascanion vetustus Baird and Girard,
Cat. N. A. Reptiles in Smiths. Inst., pt. 1, 1853, p. 97.
Type Locaniry—San Jose, Santa Clara County, California (so
restricted by Van Denburgh, Oce. Papers Calif. Acad. Sei., 5, 1897
p. 183).
Synonyms.—Bascanion constrictor flaviventris; Bascanion con-
strictor vetustum; Zamenis constrictor flaviventris; Bascanion con-
strictor; Zamenis constrictor; [2] Bascanium flagelliforme testaceum,
part.
Common Names.—Blue Racer; California Black Snake; Black
Chaser; Yellow Coachwhip Snake, part; Yellow-bellied Black Snake ;
Green Racer.
Rance.—Throughout nearly all of the state, except on the south-
eastern deserts. Oceurs south, east of the Sierra Nevada, at least to
Honey Lake, Lassen County (Yarrow and Henshaw, Ann. Rep. U.S.
Engineers, 1878, p. 1636); in central California, east to Yosemite
Valley, Mariposa County (Van Denburgh, Oce. Papers Calif. Acad.
190 University of California Publications in Zoology [Vou.17
Sci., 5, 1897, p. 186), and to Kernville, Kern County (Yarrow, U. S.
Nat. Mus. Bull., 24, 1882, pp. 16, 110) ; in southern California, east to
Fort Tejon, Kern County (Yarrow, loc. cit.), to San Bernardino (Van
Denburgh, loc. cit., p. 185), and south to Agua Caliente, 3400 feet
altitude, in San Diego County (Van Denburgh, loc. cit.). Occupies
the Lower and Upper Sonoran and Transition life-zones. Inhabits
grasslands and wet meadows. (See fig. 13.)
Coluber flagellum frenatus (Stejneger)
Red Racer
OriGinaL Description.—Bascanion flagellum frenatwm Stejneger,
N. Amer. Fauna, 7, May 31, 1893, pp. 208-209.
Type Locairy.
Mountain Spring, edge of Colorado Desert,
eastern San Diego County, California.
Synonyms.—Zamensis flagellum flagellum; Zamenis flagellum;
Zamenis flagelliformis frenatus; Bascanion flagellum frenatum;
Bascanium flageluforme; Bascanium flagelliforme testaceum, part;
Bascanium testaceum; [?] Bascanium flagelliforme piceum; Herpe-
todryas flavigularis ; Drymobius testaceus.
Common Names.—Western Whip Snake; Yellow Coach-whip
Snake, part; [?] Arizona Coach-whip Snake; Coppery Whip Snake.
Rance.—Throughout the desert and coast districts of the southern
half of the state, including also the southern San Joaquin Valley.
Has been taken northwest to Yosemite Valley, Mariposa County (Van
Denburgh, Proe. Calif. Acad. Sei., ser. 2, 5, 1895, p. 148, footnote), to
Fresno (Yarrow, U.S. Nat. Mus. Bull., 24, 1882, pp. 17, 112), and to
Carrizo Plain, San Luis Obispo County (Mus. Vert. Zool.) ; east of
the Sierras, north to Deep Spring Valley, Inyo County (Stejneger,
N. Amer. Fauna, 7, 1893, p. 209). Occurs chiefly within the Lower
Sonoran life-zone, extending rarely into Upper Sonoran and doubt-
fully into Transition. Inhabits open washes, plains, and hillslopes.
(See fig. 13.)
Coluber lateralis (Hallowell)
California Striped Racer
OrieinaL Drescription.—Leptophis lateralis Hallowell, Proc. Acad.
Nat. Sei. Phila., 6, 1853, p. 237.
Type Locaurry.—California.
1917] Grinnell-Camp: Amphibians and Reptiles of California 191
Synonyms.—Zamenis lateralis; Bascanion laterale; Bascanium
taeniatum laterale; Bascanion taeniatus, part; Drymobius lateralis.
Common NameEs.—Striped Racer, part; Hallowell’s Coach-whip
Snake; Banded Racer; Few-striped Whip Snake; Striped-side Whip
Snake.
= - r
i
LAS SES SSE
SEBS Gg if Ye
(
Coluber constrictor vetustus
Coluber flagellum frenatus
Coluber taeniatus
> beO
Coluber lateralis
Ranges of Coluber constrictor vetustus
and Coluber flagellum frenatus
outlined
Ba ONS
ee N
bs
AN
hel aes
Kon |
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA
Se
Fig. 13. Distribution of Racers (Coluber) in California.
RANGE.—Southern and west-central portions of the state, chiefly
west of the desert divides. Has been taken north to 7 miles west of
Cazadero, in Sonoma County (Mus. Vert. Zool.), to Mount Saint
Helena, in Lake County (Van Denburgh, Oce. Papers Calif. Acad.
Sci., 5, 1897, p. 190), and to Baird, Shasta County (Cope, Ann. Rep.
192 University of California Publications in Zoology (Vou. 17
U.S. Nat. Mus., 1898 [1900], p. 808) ; northeast to 5 miles northeast
of Coulterville, 3200 feet altitude, in Mariposa County (Mus. Vert.
Zool.) ; east to Fay Creek near Weldon (Mus. Vert. Zool.) and Fort
Tejon (Yarrow and Henshaw, Ann. Rep. U. 8. Engineers, 1878, p.
1637), in Kern County; to Sierra Madre, 2000 feet altitude, Los
Angeles County (Mus. Vert. Zool.) ; to Cabezon, Strawberry Valley,
6000 feet altitude, and Kenworthy, 4500 feet altitude, in Riverside
County (Atsatt, Univ. Calif. Publ. Zool., 12, 1913, p. 48); and to
Dulzura, San Diego County (Mus. Vert. Zool.). Oceupies chiefly the
Upper Sonoran life-zone, but extends locally into Lower Sonoran and
Transition. Inhabits, as a rule, chaparral. (See fig. 13.)
Coluber taeniatus (Hallowell)
Nevada Striped Racer
OrictnAL Description.—Leptophis taeniata Hallowell, Proe. Acad.
Nat. Sei. Phila., 6, 1852, p. 181.
Type Locauiry.—New Mexico.
Synonyms.—Zamenis taeniatus; Masticophis taeniatus; Basca-
nion taeniatum; Drymobius taeniatus.
Common NameEs.—Many-striped Whip Snake ; Striped Racer, part ;
Pacific Coach-whip Snake; Striped Whip Snake.
RANGE.—Chiefly east of the Sierran divides and north of the
Mohave desert. Recorded west at the north to Baird and Canoe Creek,
in Shasta County (Stejneger, N. Amer. Fauna, 7, 1893, p. 210), and
southeast through the Inyo region to Maturango Spring, Argus Range,
Inyo County (Stejneger, loc. cit.). Southernmost station, Walker
Basin, Kern County (Yarrow and Henshaw, Ann. Rep. U. S. Engi-
neers, 1878, p. 1637). Oceurs chiefly within the Upper Sonoran life-
zone. Inhabits sagebrush. (See fig. 13.)
Arizona elegans Kennicott
Faded Snake
ORIGINAL DeEscripTion.—Arizona elegans Kennicott, in Baird, U.
S. Mex. Bound. Survey, 2, 1859, pt. 2, Reptiles, pp. 18-19, pl. 13.
Type Locauiry.—Rio Grande.
Synonyms.—Rhinechis elegans; Coluber arizonae.
Common Name.—Smooth-sealed Coluber.
Rance.—The extreme southern portion of the state. All definite
1917] Grinnell-Camp: Amphibians and Reptiles of California 193
stations of occurrence known to us are as follows: Fresno (U.S. Nat.
Mus.) ; Alhambra, Los Angeles County (U.S. Nat. Mus.) ; near On-
tario, San Bernardino County (Van Denburgh, Oce. Papers Calif.
Acad. Sci., 5, 1897, p. 194) ; Riverside (Van Denburgh, Proe. Calif.
Acad. Sci., ser. 4, 3, 1912, p. 150); San Jacinto, Riverside County
(Van Denburgh, 1897, loc. cit.) ; Warner’s Ranch (Boulenger, Cat.
Snakes Brit. Mus., 2, 1894, p. 66), between Carlsbad and Oceanside
(Van Denburgh, 1897, loc. cit.), Vallecito (Mus. Vert. Zool.), and
Pacifie Beach (Cope, Ann. Rep. U. 8. Nat. Mus., 1898 [1900], p. 865),
in San Diego County. Occurs chiefly within the Lower Sonoran life-
zone. Inhabits open flat country.
Pituophis catenifer catenifer (Blainville)
Western Gopher Snake
OriaginaAL Descriprion.—Coluber catenifer Blainville, Nouy. Ann.
Mus. Hist. Nat., 4, 1835, pp. [58-59] 290-291, pl. 26, figs. 2, 2a, 2b.
Typr Locauiry.—California.
Synonyms.—Pituophis annectens Baird and Girard, Cat. N. A.
Reptiles in Smiths. Inst., pt. 1, 18538, p. 72 (type from San Diego) ;
Pityophis Heermanni Hallowell, Proe. Acad. Nat. Sei. Phila., 6, 1853,
p. 286 (type from mines in vicinity of Cosumnes River [in Eldorado
or Amador County]); Petyophis vertebralis; Pityophis sayv bellona,
part; Pityophis sayt; Pitwophis bellona, part ; Pituophis melanoleucus ;
Pituophis melanoleucus var. catenifer.
Common Names.—Pacifie Bull Snake; Bellona Bull Snake; West-
ern Bull Snake, part; Pacific Pine Snake; Gopher Snake, part; Say’s
Pine Snake; Yellow Gopher Snake.
Rance.—The whole length of the state west of the desert divides,
but chiefly east of the coast redwood belt. Also found on Santa Cruz
and Santa Catalina islands (Van Denburgh, Proe. Calif. Acad. Sei.,
ser. 4, 4, 1914, pp. 133, 136, 188). Oceupies Lower and Upper Sonoran
and Transition life-zones. Shows no particular restriction as regards
habitat, though certainly not aquatie.
Pituophis catenifer deserticola Stejneger
Desert Gopher Snake
OriatInaL Description.—Pituophis catenifer deserticola Stejneger,
N. Amer. Fauna, 7, May 31, 1893, pp. 206-208.
194 University of California Publications in Zoology [Vou.17
Type Locaurry.—Great Basin and southwestern deserts [= east
slope of Beaverdam Mountains, southwestern Utah (U. 8. Nat. Mus.,
no. 18070) |.
Synonyms.—Pityophis sayt bellona, part; Pityophis catenfer,
part; Pityophis bellona, part.
Common Names.—Western Bull Snake, part; Southern Bull
Snake; Arizona Bull Snake; Gopher Snake, part.
RanGe.—East of the desert and Great Basin divides, the whole
length of the state. Oceupies the Lower and Upper Sonoran life-zones.
Inhabits nearly all types of arid environment.
Subfamily BorginaE
Tantilla eiseni Stejneger
California Tantilla
OriGInAL DescripTION.—Tantilla eiseni Stejneger, Proe. U. 8. Nat.
Mus., 18, April 16, 1896, pp. 117-118.
Typr Locauiry.—F resno, California.
Synonym.—Tantilla nigriceps.
Common Names.—Fisen’s Black-headed Snake; Black-headed
Tantilla.
Rance.—The southern portion of the state. Only three record
stations to date: Fresno, Fresno County (as above); near Mohave,
Kern County (one specimen in Southwest Museum at Los Angeles,
fide C. L. Camp), and near Los Angeles (Riithling, Copeia, no. 15,
February 20, 1915). Seems to belong to the Lower Sonoran life-zone.
(See fig. 12.)
Superfamily VIPEROIDEAE
Family CROTALIDAE
Crotalus oreganus Holbrook
Pacifie Rattlesnake
OrieinaL Description.‘ Crotalus oreganus Holbrook, N. Amer.
Herpetology, Ist ed., 4, 1840, p. 115, pl. 29 [= 24]’’ (see Gill, Science,
ser. 2, 17, 1903, pp. 910-912).
Type Locaurry.—Columbia River.
Synonyms.—Crotalus lucifer Baird and Girard, Proe. Acad. Nat.
Sei. Phila., 6, 1852, p. 177 (types from Oregon and California) ;
1917] Grinnell-Camp: Amphibians and Reptiles of California 195
Crotalus confluentus; Crotalus lecontet, part; [?] Crotalus ruber,
part ; Crotalus adamanteus var. lucifer; Crotalus oregonus var. lucifer ;
Crotalus confluentus lucifer; Caudisona lucifer; Crotalus Hallowelli
Cooper, Amer. Nat., 3, 1870, p. 187 (range on southern coast slope of
a
Crotalus oreganus
Crotalus tigris
Crotalus mitchellii
Crotalus atrox
Crotalus exsul |
opde@40
Crotalus cerastes
Range of Crotalus oreganus outlined |
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNLA
So Se
Fig. 14. Distribution of Rattlesnakes (Crotalus) in California.
California north to 140 miles north of 30° 30’—‘‘nom. prov.’’) ; [?]
Crotalus adamanteus atrox, part.
Common Names.—Black Rattlesnake; California Rattlesnake ;
Arizona Diamond Rattlesnake, part; Confluent Rattlesnake; Oregon
Rattlesnake ; Missouri Rattlesnake ; Hallowell’s Rattlesnake ; Southern
Rattlesnake.
196 University of California Publications in Zoology [Vou.17
Rance—Throughout the state chiefly west and north of the Colo-
rado and Mohave deserts. Extends to an altitude of 8600 feet on the
central Sierra Nevada. Has been found on the southeast to Charlotte
Creek, 8500 feet altitude, Fresno County (Mus. Vert. Zool.) ; to Walker
Pass (Mus. Vert. Zool.) and Mohave (Meek, Field Columb. Mus., zool.
ser., 7, 1905 [1906], p. 17), in Kern County; to Pine Flats, 5500 feet
altitude, San Gabriel Mountains, Los Angeles County (Grinnell and
Grinnell, Throop Inst. Bull., 35, 1907, p. 53); to Doble, 7000 feet
altitude, San Bernardino Mountains, San Bernardino County (Grin-
nell, Univ. Calif. Publ. Zool., 5, 1908, p. 53); to Banning, Tahquitz
Valley, 8000 feet altitude, and Santa Rosa Peak, in Riverside County
(Atsatt, Univ. Calif. Publ. Zool., 12, 1913, p. 45) ; and to summit of
Coast Range, near Mexican boundary, in San Diego County (Cope,
Ann. Rep. U.S. Nat. Mus., 1898 [1900], p. 1179). Also found on the
Providence Mountains, eastern San Bernardino County (U. 8. Nat.
Mus.), and on Santa Catalina Island (Yarrow, U. S. Nat. Mus. Bull.,
24, 1882, p. 76). Oceupies all life-zones from Lower Sonoran to
Canadian. Inhabits almost all types of environment, though ap-
parently commonest on chaparral slopes and in open country where
eround-squirrel burrows abound. (See fig. 14.)
Crotalus atrox Baird and Girard
Texas Rattlesnake
OrieginaL Descrietion.—Crotalus atrox Baird and Girard, Cat.
N. A. Reptiles in Smiths. Inst., pt. 1, 1853, pp. 5-6.
Type Locauiry.—Indianola or San Pedro, Texas.
Synonyms.—Crotalus adamanteus atrox, part; Crotalus lecontet,
part.
Common Names.—Western Diamond Rattlesnake, part; Fierce
Rattlesnake; Arizona Diamond Rattlesnake, part.
Ranee.—Colorado Desert near Mexican boundary. Recorded from
Fort Yuma (Yarrow, U.S. Nat. Mus. Bull., 24, 1882, pp. 12, 75) and
Laguna Station, New River (Cope, Ann. Rep. U. 8. Nat. Mus., 1898
[1900], p. 1167) ; both stations in Imperial County. Life-zone, Lower
Sonoran. (See fig. 14.)
Crotalus exsul Garman
Red Rattlesnake
OrigInaL Description —Crotalus exsul Garman, Mem. Mus. Comp.
Zool., 8, no. 3, June, 1883, pp. 114-115, 174. [Dr. Thomas Barbour
1917] Grinnell-Camp: Amphibians and Reptiles of California 197
has examined Garman’s type and finds it to belong to the species which
has currently borne the name ruber. |
Type Locaurry.—Cedros Island, Lower California.
Synonyms.—Crotalus atrox, part; Crotalus adamanteus ruber
Cope, Proce. U. 8S. Nat. Mus., 14, 1891, pp. 690-691 (type locality
unknown) ; Crotalus atrox ruber; Crotalus ruber.
Common Names.—Red Diamond Rattlesnake; Western Diamond
_ Rattlesnake, part.
Rance.—Extreme southwestern corner of the state. Oceurs north
to Reche Caton (Camp, MS) and Cabezon (Atsatt, Univ. Calif. Publ.
Zool., 12, 1913, p. 44), in Riverside County ; east to Dos Palmos Spring,
3500 feet altitude, Santa Rosa Mountains, Riverside County (Atsatt,
loc. cit.), and to Mountain Spring, in San Diego County (Cope, Ann.
Rep. U. 8. Nat. Mus., 1898 [1900], p. 1169) ; west to Deluz (Van
Denburgh, Oce. Papers Calif. Acad. Sei., 5, 1897, p. 228), to Twin
Oaks (Stejneger, Ann. Rep. U. S. Nat. Mus., 1893, p. 440), and to
El Nido P. O. (Cope, loc. cit.), in San Diego County. Oceupies the
Upper Sonoran life-zone. Inhabits chiefly chaparral-covered and
rocky hillslopes. (See fig. 14.)
Crotalus tigris Kennicott
Tiger Rattlesnake
OrigiNAL Description.—Crotalus tigris Kennicott, U. S. Mex.
Bound. Surv., 2, 1859, pt. 2, Reptiles, p. 14, pl. 4.
Type Locauiry.—Sierra Verde and Pozo Verde [Sierra del
Pozo Verde, Arizona: Stejneger, N. Amer. Fauna, 7, 1893, p. 214].
RanGeE.—South-central portion of the state east of the Sierra
Nevada, chiefly in desert ranges of Inyo County. Recorded north-
west to Beveridge Canon, 8000 feet altitude, Inyo Mountains (Meek,
Field Columb. Mus., zool. ser., 7, 1905 [1906], p. 16) ; west to Inde-
pendence Creek and Coso Valley, and south to Slate Range, 3100 feet
altitude (Stejneger, loc. cit.). Life-zone, chiefly Upper Sonoran. In-
habits rocky situations. (See fig. 14.)
Crotalus mitchellii (Cope)
Pallid Rattlesnake
OrIGInAL DeEscripTion.—Caudisona mitchellii Cope, Proc. Acad.
Nat. Sci. Phila., 1861, pp. 293-294.
Type Locatiry.—Cape San Lueas, Lower California.
198 University of California Publications in Zoology [Vou.17
Synonyms.—Crotalus pyrrhus; [2] Crotalus lecontei, part.
Common Names.—Bleached Rattlesnake; White Rattlesnake.
Rance.—The Colorado and Mohave deserts. Has been taken north-
west to Fairmont, northern Los Angeles County (Grinnell and Grin-
nell, Throop Inst. Bull., 35, 1907, pp. 59-60) ; northeast to 14 miles
northeast of Blythe Junction, San Bernardino County (Camp, Univ.
Calif. Publ. Zool., 12, 1916, pp. 533-534) ; west to 5 miles southwest
of Banning, in Riverside County (Mus. Vert. Zool.), to Asbestos
Spring, Santa Rosa Mountains, Riverside County (Atsatt, Univ. Calif.
Publ. Zool., 12, 19138, p. 44), and to Mountain Spring, San Diego
County (Van Denburgh, Proce., Calif. Acad. Sci., ser. 2, 4, 1894, pp.
450-455). Oceupies the Lower Sonoran life-zone, extending locally
into Upper Sonoran. Inhabits nearly all types of arid environment.
(See fig. 14.)
Crotalus cerastes Hallowell
Sidewinder
OriGINAL DeEscripTion.—Crotalus cerastes Hallowell, Proe. Acad.
Nat. Sei. Phila., 7, 1854, pp. 95-96.
Type Locatiry.—Borders of the Mohave River and in the desert
of the Mohave.
Common Name.—Horned Rattlesnake.
RancGe.—Colorado and Mohave deserts; also north into the Inyo
region. Northernmost stations are: Mesquite Valley (Meek, Field
Columb. Mus., zool. ser., 7, 1905 [1906], p. 18) and Lone Pine (Stej-
neger, N. Amer. Fauna, 7, 1893, p. 218), in Inyo County ; westernmost
stations are: Oro Grande, in San Bernardino County (Meek, loc. cit.),
Torres [Toro], west of Mecea, in Riverside County (Mus. Vert. Zool.),
and Coyote Wells, in Imperial County (Cope, Ann. Rep. U. S. Nat.
Mus., 1898 [1900], p. 1199). Occupies the Lower Sonoran life-zone.
Restricted to sand-dune areas and level tracts of loose sandy soil. (See
fig. 14.)
Order TESTUDINATA
Suborder ATHECAE
Family DERMOCHELIDAE
Dermochelys schlegelii (Garman)
Pacific Leatherback Turtle
OrigiInaL Description.—Sphargis schlegelii Garman, U. 8. Nat.
Mus. Bull., 25, 1884, pp. 294-295 (see also p. 303).
1917] Grinnell-Camp: Amphibians and Reptiles of California 199
Tyre Locauiry.—Tropiecal Pacific and Indian oceans.
Rance.—Ocean off southern coast. Three known instances of
capture: Santa Barbara (two specimens), and off Point Loma, San
Diego County (one specimen) (Van Denburgh, Proe. Calif. Acad.
Sci., ser. 3, zool., 4, 1905, pp. 51-60, pls. 9-11).
Suborder LAMINIFERA
Family TESTUDINIDAE
Subfamily Emyprmvar
Clemmys marmorata (Baird and Girard)
Pacific Mud Turtle
ORIGINAL DEscriIPTION.—E mys marmorata Baird and Girard, Proe.
Acad. Nat. Sci. Phila., 6, 1852, p. 177.
Type Locauiry.—Puget Sound.
Synonyms.—Emys nigra Hallowell, Proe. Acad. Nat. Sei. Phila.,
7, 1854, pp. 91-92 (type from ‘‘Posa Creek, Lower California’’
[= Poso Creek, Kern County?]|); Actinemys marmorata; Clemmys
Wosnessenskyt Strauch, Mem. Acad. Imper. Sei. St. Petersb., ser. 7, 5,
no. 7, 1862, pp. 114-117, pl. opp. p. 196 (type from Rio Sacramento,
California) ; Chelopus marmoratus.
Common Names.—Western Pond Turtle; Pacifie Terrapin; Cali-
fornia Terrapin; California Mud Turtle; Western Terrapin; Water
Turtle.
Ranee.—Most of the streams on the Pacifie watershed, the whole
length of the state. Has been reported east to Pit River [in Shasta
County?] (Townsend, Proc. U. S. Nat. Mus., 10, 1887, p. 237); to
6 miles east of Coulterville, 2800 feet altitude, in Mariposa County
(Mus. Vert. Zool.) ; to South Fork of Kern River, 25 miles above
Kernville, in Kern County (Stejneger, N. Amer. Fauna, 7, 1893, p.
162) ; and to Mohave River, in San Bernardino County (Cooper, Amer.
Nat., 3, 1870, p. 189); and south to San Diego, San Diego County
(Yarrow, U.S. Nat. Mus. Bull., 24, 1882, pp. 7, 36).
Subfamily TesTuDININAE
Testudo agassizli (Cooper)
Desert Tortoise
OriGgInaL DeEscripTion.—NXerobates agassizii Cooper, Proe. Calif.
Acad. Sei., 2, 1863, pp. 120-121.
200 University of California Publications in Zoology [Vou.17
Type Locaurry.—Mountains of California near Fort Mohave
[=‘‘Salado Valley,’’ on Mohave Desert: True, Proc. U. S. Nat. Mus.,
4, 1881 (1882), p. 447].
Synonyms.—Gopherus agassizii; Xerobates berlandiert.
Common Names.—Agassiz’s Gopher; Western Gopher; Agassiz’s
Tortoise; Agassiz’s Land Tortoise.
RanGcE.—Chiefly the Mohave Desert. Reported north as far as
Crater Summit (Van Denburgh, Oee. Papers Calif. Acad. Sci., 5, 1897,
p- 37) and Leach Point Valley (Stejneger, N. Amer. Fauna, 7, 1893,
p- 162), in northern San Bernardino County; west to one-half mile
east of Mohave, Kern County, and to 3 miles south of Palmdale, Los
Angeles County (Camp, Univ. Calf. Publ. Zool., 12, 1916, p. 513) ;
south to Cottonwood Mountains, Riverside County (Camp, loc. cit.),
and, possibly, as far as Fort Yuma (True, loc. cit.), Imperial County.
Restricted to the Lower Sonoran life-zone. Inhabits, as a rule, flat
gravelly or sandy tracts, but found also on rocky hills. (See fig. 9.)
Family KINOSTERNIDAE
Kinosternon sonoriense LeConte
Arizona Mud Turtle
OrtqinaL Descrietion.—Kinosternum sonoriense LeConte, Proce.
Acad. Nat. Sci. Phila., 7, 1854, p. 184.
Typp Locauiry—Tueson, Sonora [= Arizona].
Synonyms.—Cinosternum flavescens; Platythyra flavescens.
Common Name.—Yellow Mud Turtle.
Rance—Lower Colorado River. Two definite stations of oceur-
rence: California side of the Colorado River opposite Yuma (Van
Denburgh, Proe. Calif. Acad. Sei., ser. 4, 3, 1913, p. 396 [see also
Cooper, in Cronise, Nat. Wealth Calif., 1868, p. 481] ) ; and Palo Verde
(Mus. Vert. Zool.).
Transmitted November 21, 1916.
1917] Grinnell-Camp: Amphibians and Reptiles of California 201
A
Ablabes punctatus, 184.
purpureocauda, 183.
Actinemys marmorata, 199.
Agama coronatum, 164.
Douglassii, 164.
Amblystoma californicum, 138.
paroticum, 139.
rubrum, 130.
tenebrosum, 139.
Ambyostoma californiense, 138.
Ambystoma ensatum, 131, 139.
ingens, 138.
macrodactylum, 131, 138.
mavortium, 138.
mavortium californiense, 138.
paroticum, 131, 139.
punctatum, 134.
punctulatum, 134.
tigrinum, 131, 138.
tigrinum californiense, 138.
Ambystomidae, 138.
Amphibia, 130.
Anaides ferreus, 135.
jécanus, 135.
lugubris, 134, 135.
Anaides, sad-colored, 130, 134.
Aneides ferreus, 133, 135.
iécanus, 133, 135.
lugubris, 130.
lugubris farallonensis, 133, 135.
lugubris lugubris, 133, 134.
Anguidae, 166.
Anniella nigra, 170, 171.
pulchra, 170, 171.
pulchra nigra, 170.
pulechra pulchra, 170.
texana, 170.
Anniellidae, 170.
Anolis carolinensis, 157.
cooperi, 157.
principalis, 157.
Anota calidiarum, 165.
M ’Callii, 166.
platyrhina, 165.
Arizona elegans, 192.
Armor-bearer, Tiger, 172.
Ascaphus truei, 139.
Athecae, 198.
Autodax iécanus, 135.
lugubris, 134, 135.
lugubris farallonensis, 135.
Autodax, spotted, 134.
Axolotl, California, 138.
B
Bascanion constrictor, 189.
constrictor flaviventris, 189.
constrictor vetustum, 189.
INDEX
flagellum frenatum, 190.
laterale, 191.
taeniatum, 192.
taeniatus, 191.
vetustus, 189.
Bascanium flagelliforme, 190.
flagelliforme piceum, 190.
flagelliforme testaceum, 189, 190.
taeniatum laterale, 191.
testaceum, 190.
Batrachoseps attenuatus, 136, 137.
major, 136, 137.
nigriventris, 137.
pacificus, 136, 137.
Batrachosoma coronatum, 165.
Bellophis zonatus, 185.
Bell-toad, American, 139.
Boa, California, 177.
Rosy, 177.
Rubber, 177, 178.
Boidae, 177.
Boiginae, 194.
Boinae, 177.
Bufo alvarius, 141, 144.
beldingi, 144.
boreas, 142, 143.
boreas boreas,.141, 143.
boreas halophilus, 141, 142, 143.
boreas nelsoni, 141, 142, 143.
canorus, 141, 143.
chilensis, 142.
cognatus, 140.
cognatus californicus, 141.
cognatus cognatus, 140, 141.
columbiensis, 142, 143.
columbiensis halophilus, 142.
dorsalis, 142.
halophila, 142.
halophilus, 143.
lentiginosus americanus, 142.
lentiginosus woodhousii, 142.
microseaphus, 143.
punctatus, 141, 144.
woodhousii, 141, 142.
Bufonidae, 140.
Bull-frog, 149.
Cc
Callisaurus draconoides, 151.
draconoides ventralis, 151.
dracontoides gabbii, 151.
ventralis ventralis, 151.
Caudata, 130.
Caudisona lucifer, 195.
mitchellii, 197.
Charina bottae, 177, 178.
brachyops, 178.
plumbea, 178.
Chaser, Black, 189.
Chelopus marmoratus, 199.
202 University of California Publications in Zoology
Chilomeniscus cinctus, 182, 185.
ephippicus, 182.
Chionactis occipitalis, 182.
occipitalis annulatus, 182.
Chondrotus lugubris, 139.
tenebrosus, 139.
Chuckwalla, 153.
Cinosternum flavescens, 200.
Clemmys marmorata, 199.
Wosnessenskyi, 199.
Cnemidophorus gracilis, 172.
grahamii, 174.
grahamii stejnegerii, 174.
hyperythrus, 175.
hyperythrus beldingi, 173, 175.
sexlineatus var. bocourtii, 172.
sexlineatus var. tesselatus, 172.
stejnegeri, 174.
tessellatus tessellatus, 172, 174.
tessellatus tigris, 172, 174.
tigris, 172.
tigris mundus, 172, 173.
tigris stejnegeri, 172, 173, 174.
tigris tigris, 172, 173.
tigris undulatus, 174.
undulatus, 173.
Coleonyx variegatus, 149, 167.
Coluber arizonae, 192.
catenifer, 193.
constrictor vetustus, 189, 191.
flagellum frenatus, 190, 191.
infernalis, 181.
lateralis, 190, 191.
(Ophis) Californiae, 187.
parietalis, 181.
punctatus, 184.
taeniatus, 191, 192.
(Zacholus) zonatus, 184.
Coluber, Smooth-sealed, 192.
Colubridae, 179.
Colubrinae, 182.
Colubroideae, 179.
Contia mitis, 183, 185.
Coronella amabilis, 184.
balteata, 186.
boylii, 186.
Californiae, 187.
getula, 186, 187.
getulus var. californica, 187.
multifasciata, 185.
pyromelanus zonata, 185.
zonata, 185.
Costata, 139.
Crotalidae, 194.
Crotalus adamanteus atrox, 195, 196.
adamanteus var. lucifer, 195.
adamanteus ruber, 197.
atrox, 195, 196, 197.
atrox ruber, 197.
cerastes, 195, 198.
confluentus, 195.
confluentus lucifer, 195.
exsul, 195, 196.
Hallowelli, 195.
lecontei, 195, 196, 198.
lucifer, 194.
mitchellii, 195, 197.
oreganus, 194, 195.
oregonus var. lucifer, 195.
pyrrhus, 198.
ruber, 195, 197.
tigris, 195, 197.
Crotaphytus baileyi, 153.
collaris, 153.
collaris baileyi, 153.
dorsalis, 150.
fasciatus, 152.
Gambelii, 152.
silus, 152.
wislizenii, 152.
Cynops torosus, 130.
D
Dermochelidae, 198.
Dermochelys schlegelii, 198.
Diadophis amabilis, 184, 185.
amabilis pulchellus, 184.
pulchellus, 184.
punctatus, 184.
punctatus amabilis, 184.
punctatus pulchellus, 184.
Dicamptodon ensatus, 139.
Diemictylus torosus, 130.
Dipso-saurus dorsalis, 150.
Discoglossidae, 139.
Doliosaurus me’ealli, 166.
platyrhinos, 165.
Dragon, Spotted-tail, 151.
Drymobius lateralis, 191.
taeniatus, 192.
testaceus, 190.
E
Elgaria formosa, 169.
scincicauda, 167.
Emydinae, 199.
Emys marmorata, 199.
nigra, 199.
Ensatina Eschscholtzii, 132.
Epirhexis longipes, 149.
Eublepharidae, 149.
Eublepharis variegatus, 150.
Eumeces gilberti, 175.
hallowellii, 176.
quadrilineatu[s], 175.
skiltonianus, 176.
skiltonianus var. amblygrammus,
176.
skiltonianus var. brevipes, 176.
Euphryne obesus, 153.
Eutaenia coneinna, 181.
couchii, 180, 181.
elegans, 179.
elegans brunnea, 180.
elegans couchii, 180, 181.
elegans infernalis, 180.
elegans lineolata, 180.
[ Vou. 17
1917] Grinnell-Camp: Amphibians and Reptiles of California 203
elegans marciana, 179. Gopher, Agassiz’s, 200.
elegans ordinoides, 179. Western, 200.
elegans vagrans, 180. Gopherus agassizii, 200.
Hammondii, 180, 181. Greeny, 145.
imperialis, 181.
infernalis, 179, 180. H
infernalis vidua, 179.
leptocephala, 179.
marciana, 181.
ordinoides, 179.
proxima, 181.
Hemidactylium pacificum, 136.
Herpetodryas flavigularis, 190.
Heterotriton ingens, 138.
Homalosoma mite, 183.
Homalosaurus ventralis, 151.
sirtalis, 181. Horned-toad, Blainyille, 164.
sirtalis dorsalis, 181. Galifornia, 165.
sittalis leptocephala, 179. Mesernsl65 ae
sirtalis obseura, 181. Flat-tailed 166.
sirtalis parietalis, 181. pienie 164.
sirtalis pickeringii, 181. Hyla affinis 145.
sirtalis tetrataenia, 181. amenicalor 145.
Eutainia atrata, 179. cadaverina, 145.
elegans, 180. copii, 145.
Marciana, 179. nebulosa, 145.
sirtalis elegans, 179. regilla, 144, 145.
vagrans, 180. regilla scapularis, 145,
scapularis, 145.
F scapularis hypochondriaca, 145.
’ versicolor, 145
Frog, Boyle’s, 146. VOSS) Teh 7
California Red-legged, 148. Hyla, Cadaverous, 145.
Cape San Lucas, 145.
Cope’s, 145.
Pacific, 145.
Hylidae, 144.
Hylinae, 144.
California Yellow-legged, 146.
Drayton’s, 149.
Leconte’s, 149.
Leopard, 149.
Long-footed, 149.
Nevada Spotted, 148. Hypsiglena ochrorhynchus, 178, 188.
Oregon Red-legged, 148.
Pacific, 146. I
Rocky Mountain, 149. Iguana, Desert, 150.
Sand-colored Tree, 145. Iguanidae, 150.
Salt-marsh, 142. Iguaninae, 150.
Sierra Madre Yellow-legged, 147.
Sierra Nevada Yellow-legged, 146. J
Thick-skinned, 146. Jack, Swift, 172.
Western, 146.
Western Spotted, 148. K
Western Wood, 149. Kinosternidae, 200.
Kinosternon sonoriense, 200.
G Kinosternum sonoriense, 200.
Gecko, Banded, 149.
Variegated, 150. L
Gerrhonotus Burnettii, 169. Laminifera, 199.
coeruleus, 167, 169. Lampropeltis boylii, 186.
grandis, 169. boylii var. conjuncta, 187.
multicarinatus, 167, 168, 169. californiae, 187.
multicarinatus palmerii, 169. conjuncta, 187.
palmeri, 167, 168. pyromelana multicincta, 184.
scincicauda, 168. zonata, 185.
scincicauda ignavus, 168. Lamprosoma annulatum, 182.
scincicauda palmeri, 168. episcopum, 183.
scincicauda scincicauda, 166, 167. occipitale, 182.
scincicauda webbii, 167, 168. Leptophis lateralis, 190.
scincicaudus, 169. taeniata, 192.
webbii, 168. Leptotyphlopidae, 176.
Glauconia humilis, 177. Leptotyphlops humilis, 176.
204 University of California Publications in Zoology
Lichanura myriolepis, 177.
oreutti, 177.
roseofusea, 177, 178.
simplex, 177.
trivirgata, 177.
Linguata, 140.
Litoria occidentalis, 145.
Lizard, Alderman, 154.
Alligator, 159.
Bailey Collared, 153.
Bailey Leopard, 153.
Banded, 152.
Beautiful, 151.
Belding Orange-throated, 175.
Black-bellied, 137.
Black Footless, 170.
Blainville’s Horned, 164, 165.
Blue-bellied, 161.
Blue-tailed, 176.
Broad-nosed Barrel, 166.
Brown-shouldered, 156.
Burnett’s Alligator, 169.
Burnett’s Keeled, 169.
California Alligator, 166.
California Horned, 164, 165.
California Whip-tailed, 173.
Cape Striped, 175.
Clark’s, 162.
Clark’s Alligator, 162.
Coast Alligator, 169.
Colorado Desert, 150.
Cooper’s Green, 157.
Cope’s, 171.
Cope’s Desert, 151.
Crested, 150.
Jrowned Horned, 164, 165,
Desert Brown-shouldered, 155.
Desert Horned, 165.
Desert Night, 171.
Desert Whip-tailed, 172.
Dusky Rough-sealed, 162, 163.
Dusky Sealy, 163.
Fat, 154.
Fat Toad, 154.
Fence, 157, 160, 162.
Flat-tailed Horned, 166.
Graceful, 157.
Graham’s Striped, 174.
Great Basin Leopard, 152.
Mountain Alligator, 169.
New Mexican Alligator, 157.
Northern Brown-shouldered, 154.
Ocellated Desert, 151.
Ocellated Sand, 150.
Pacific, 136.
Pacific Blue-bellied, 159.
Pacific Horned, 164.
Pigmy Horned, 164.
Red-spotted Desert, 151.
Rivers’s, 172.
San Diego Alligator, 168.
San Joaquin Leopard, 152.
Sealy, 162.
Schott’s, 157.
Sharp-back, 150.
Short-nosed Leopard, 152.
Sierran Alligator, 168.
Silvery Footless, 170.
Skink-tailed, 167, 169.
Slender, 137, 157.
Smooth Horned, 166.
Southern Brown-shouldered, 156.
Southern Gridiron-tailed, 151.
Spiny Alligator, 163.
Spiny-breasted Horned, 165.
Spotted Yuma, 151.
Stejneger Whip-tailed, 174.
Tenaya Blue-bellied, 160.
Tessellated, 172.
Tessellated Tiger, 174.
Thayer’s Alligator, 159, 161.
Thirsty, 150.
Tiger, 172.
Two-lined, 161.
Two-striped, 161.
Van Denburgh, 159.
Variegated, 150.
Webb’s, 168.
Western, 159.
Western Alligator, 161.
Western Fence, 159.
Wislizenius’, 152.
Worm, 170.
Xantus’s, 171.
Yellow-spotted, 133.
Zebra-tailed, 151.
M
[ Vou, 17
Masticophis taeniatus, 192.
Green-brown, 169. Molge torosa, 130.
Henshaw’s, 171. Mutabilia, 130.
Henshaw Night, 171. N
Island Blue-bellied, 162. Natricinae, 179.
Island Night, 172. Newt, California, 130.
Keel-backed, 150. Pacific Coast, 130.
Little Horned, 164. Western, 130.
Long-footed, 161. Notophthalmus torosus, 130, 131.
Many-keeled, 167, 168, 169. Nouns, Bloody, 149.
Many-ribbed, 167.
Marey’s Alligator, 157. °
McCall’s Horned, 166. Ophibolus Boylii, 186.
Mearns, 154. ealiforniae, 187.
Mountain, 157, 159. getulus boylii, 186, 187.
Great Fence, 162.
1917] Grinnell-Camp: Amphibians and Reptiles of California
getulus californiae, 187.
getulus eiseni, 187.
getulus multicinctus, 184.
pyrrhomelanus, 185.
pyrrhomelas, 185.
zonatus, 185.
Ophisaurus ventralis, 177.
Orange-throat, Belding’s, 175.
P
Pelobatidae, 140.
Phimothyra hexalepis, 189.
Phrynosoma Blainvillii, 164, 165.
blainvillii blainvillii, 164, 165.
blainvillii frontale, 165.
calidiarum, 165.
cornutum, 165.
coronatum, 164, 165.
douglassii douglassii, 164.
douglassi pygmaea, 164.
frontalis, 165.
m’eallii, 166.
modestum, 164.
platyrhinos, 165.
Pituophis annectens, 193.
bellona, 193.
eatenifer catenifer, 193.
catenifer deserticola, 193.
melanoleucus, 193. :
melanoleucus var. catenifer, 193.
Pityophis bellona, 194.
eatenifer, 194.
Heermanni, 193.
sayi, 193.
sayi bellona, 193, 194.
vertebralis, 193.
Platythyra flavescens, 200.
Plestiodon skiltonianum, 175.
Plethodon ecrassulus, 134.
croceater, 132, 133.
elongatus, 138, 134.
ensatus, 132.
eschscholtzii, 132, 133.
flavipunctatus, 133.
iécanus, 135.
intermedius, 134.
oregonensis, 132.
Plethodon, Oregon, 132.
Plethodontidae, 132.
Plethodontinae, 132.
Pleurodeles californiae, 130.
Pseudoeryx bottae, 178.
R
Racer, Banded, 191.
Blue, 189.
California Striped, 190.
Green, 189.
Nevada Striped, 192.
Red, 190.
Striped, 191, 192.
Western Yellow-bellied, 189.
Rana agilis aurora, 149.
aurora, 146, 148.
aurora aurora, 147, 148, 149.
aurora draytonii, 147, 148.
boylii, 146.
boylii boylii, 146, 147.
boylii muscosa, 147.
boylii sierrae, 146, 147.
eatesbiana, 149.
Draytonii, 148.
Lecontii, 148.
longipes, 149.
nigricans, 149.
pachyderma, 146.
pipiens, 147, 149.
pipiens brachycephala, 149.
pretiosa, 146, 148.
pretiosa luteiventris, 147, 148.
pretiosa pretiosa, 147, 148.
septentrionalis, 149.
temporaria, 149.
temporaria aurora, 149.
temporaria pretiosa, 146.
Ranidae, 146.
Raninae, 146.
Rattlesnake, Arizona Diamond,
196.
Black, 195.
Bleached, 198.
California, 195.
Confluent, 195.
Fierce, 196.
Hallowell’s, 195.
Horned, 198.
Missouri, 195.
Oregon, 195.
Pacific, 194.
Pallid, 197.
Red, 196.
Red Diamond, 197.
Southern, 195.
Texas, 196.
Tiger, 197.
Western Diamond, 196, 197.
White, 198.
Rena, California, 177.
Rena humilis, 176.
Reptilia, 149.
Rhinechis elegans, 192.
Rhinocheilus lecontei, 178, 188.
Rhinostoma occipitale, 182.
Ss
Salamander, Arboreal, 134.
Black, 135.
British Columbia, 139.
California, 138.
California Land, 134.
Capt. Beechey’s, 130.
Del Norte, 134.
Farallon, 135.
Flat-footed, 138.
Garden, 136.
Island, 136.
205
206 University of California Publications in Zoology
Long-toed, 138.
Marbled, 139.
Mount Lyell, 132.
Mournful, 134.
Oregon, 132, 139.
Rusty, 135.
Shasta, 135.
Slender, 136.
Speckled, 134.
Tiger, 138.
Vancouver’s, 139.
Warty, 130.
Yellow-spotted, 132.
Salamandra attenuata, 137.
Beecheyi, 130.
lugubris, 134.
tigrina, 138.
Salamandridae, 130.
Salamandrina attenuata, 136.
Salamandroideae, 130.
Salientia, 139.
Salvadora grahamiae, 189.
grahamiae hexalepis, 189.
hexalepis, 178, 189.
Sauria, 149.
Sauromalus ater, 153.
Scaphiopus hammondii, 140.
hammondii hammondii, 140.
Sceloporus becki, 162.
bi-seriatus, 160.
bi-seriatus var. A. azureus, 160.
bi-seriatus becki, 162.
bi-seriatus var. marmoratus, 160.
bi-seriatus var. B. variegatus, 160.
clarkii, 162.
consobrinus, 157.
consobrinus gratiosus, 157.
frontalis, 159.
graciosus, 157, 158, 159.
graciosus graciosus, 157, 158, 159.
graciosus vandenburgianus, 158,
159.
longipes, 160.
magister, 158, 162.
occidentalis, 159, 160.
occidentalis becki, 161, 162.
occidentalis bi-seriatus, 160, 161.
occidentalis occidentalis, 159, 161.
occidentalis taylori, 160, 161.
oreutti, 158, 162, 163.
spinosus, 163.
spinosus var. clarkii, 162.
undulatus bocourtii, 160.
undulatus var. bocourtii, 159.
undulatus occidentalis, 159.
undulatus thayeri, 159, 160.
undulatus undulatus, 159, 160.
vandenburgianus, 159.
Scincidae, 175.
Serpentes, 176.
Siagonodon humilis, 177.
Sidewinder, 198.
Skink, Blue-tailed, 176.
Gilbert’s, 176.
[| Vou. 17
Red-headed, 176.
Skilton’s, 176.
Western, 175.
Snake, Arizona Bull, 194.
Arizona Coach-whip, 190.
Arizona Ringed, 186.
Banded Burrowing, 182.
Banded Flat-nosed, 189.
Banded Milk, 186.
Bellona Bull, 193.
Black Garter, 179.
Black King, 187.
Blainville’s King, 187.
Blue Worm, 170.
Boyd’s Garter, 179, 180.
Boyle King, 186.
Boyle Milk, 186.
Brown, 183.
California Black, 189.
California Blind, 177.
California Garter, 181.
California King, 186, 187.
California Milk, 186, 187.
California Ring-necked, 184.
Churchill’s Garter, 182.
Coppery Whip, 190.
Coral King, 184.
Corral, 186.
Couch’s Garter, 181.
Desert, 182.
Desert Burrowing, 182.
Desert Gopher, 193.
Dusky Garter, 182.
Eisen’s Black-headed, 194.
Hisen’s King, 186.
Elegant Garter, 180.
Faded, 192.
Few-striped Whip, 191.
Gentle Brown, 183.
Glass, 177.
Gopher, 193, 194.
Graham’s Flat-nosed, 189.
Green Garter, 180.
Hallowell’s Coach-whip, 191.
Hammond’s Garter, 180, 181.
Harlequin, 186.
Horse, 182.
Lead-colored Worm, 178.
Leconte’s, 188.
Long-nosed, 188.
Many-striped Whip, 192.
Marcy Garter, 179.
Marcy’s Garter, 181.
Mohave Ringed, 182.
Narrow-headed Garter, 179.
Pacific Bull, 193.
Pacific Coach-whip, 192.
Pacifie Coast Garter, 179.
Pacific Garter, 180, 181.
Pacific Ground, 183.
Pacific Pine, 193.
Patch-nosed, 189.
Pickering’s Garter, 182.
Puget Garter, 179.
1917] Grinnell-Camp: Amphibians and Reptiles of California — 207
Purple-tailed, 183.
Red and Black Ground, 182.
Red-barred Garter, 181.
Red-bellied, 184.
Red Milk, 186.
Ring, 186.
Ringed King, 186.
Rock, 188.
Rocky Mountain Garter, 181.
Rubber, 177.
Say’s Garter, 182.
Say’s Pine, 193.
Sharp-tailed, 183.
Sheep-nosed, 177.
Single-striped Garter, 180.
Southern Bull, 194.
Spotted Night, 188.
Spotted Ring, 184.
Striped, 182.
Striped-side Whip, 191.
Striped Whip, 192.
Texas Ground, 183.
Two-headed, 178.
Wandering Garter, 180.
Water, 181.
Western Bull, 193, 194.
Western Garter, 180, 181.
Western Gopher, 193.
Western Ring-necked, 184.
Western Whip, 190.
Wood, 178.
Worm, 170, 176.
Xantus’s, 188.
Yellow-bellied Black, 189.
Yellow Coach-whip, 189, 190.
Yellow Gopher, 193.
Sonora episcopa, 183, 185.
occipitalis, 182, 185.
Spadefoot, Hammond’s, 140.
Western, 140.
Spea hammondii, 140.
stagnalis, 140.
Spea, Hammond’s, 140.
New Mexican, 140.
Spelerpes platycephalus, 132, 133.
Spelerpinae, 132.
Sphargis schlegelii, 198.
Squamata, 149.
Stenodactylus variegatus, 149.
Stenostoma humile, 177.
Swift, Common, 161.
Fence, 161.
Long-tailed, 157.
Mearns, 154.
Oreutt’s, 163.
Ornate, 157.
Pacific, 159.
Sage-brush, 157, 159.
Stansbury ’s, 156.
aineerel'o i.
Van Denburgh’s, 159.
Western, 161.
White-bellied, 157.
T
Tantilla, Black-headed, 194.
California, 194.
Tantilla eiseni, 185, 194.
nigriceps, 194.
Tapaya coronata, 165.
Douglassii, 164.
Taricha laevis, 130.
lugubris, 134.
torosa, 130.
Teiidae, 172.
‘verrapin, California, 199.
Pacific, 199.
Western, 199.
Testudinata, 198.
Testudinidae, 199.
Testudininae, 199.
Testudo agassizii, 167, 199.
Thamnophis elegans, 180, 181.
infernalis, 179, 180, 181.
leptocephala, 179.
marcianus, 179.
ordinoides elegans, 180.
ordinoides hammondii, 180, 181.
ordinoides ordinoides, 179, 180.
parietalis, 180, 181.
sirtalis parietalis, 181.
vagrans, 180.
Toad, American, 142.
Arroyo, 141.
Ashy Horned, 165.
Baird’s, 142, 143.
Broad-nosed Horned, 166.
California, 142.
Colorado, 144.
Colorado River, 144.
Common, 142.
Douglass’s Horned, 164.
Girard’s, 144.
Great Plains, 140.
MacCall’s Horncd, 166.
Nevada, 143.
Northwestern, 143.
Pacific Horned, 164.
Rocky Mountain, 142.
Small-spaded, 143.
Smooth Horned, 166.
Spotted, 144.
Woodhouse’s, 142.
Yosemite Park, 143.
Tortoise, Agassiz’s, 200.
Agassiz’s Land, 200.
Desert, 199.
Tortrix Bottae, 177.
Tree-frog, Arizona, 145.
Cope’s, 145.
Pacific, 144.
Western, 145.
Tree-toad, 145.
Triton, Cape St. Lucas, 133.
Oregon, 132.
Triton ensatus, 139.
Ermani, 130.
laevis, 130.
208 University of California Publications in Zoology
tereticauda, 130.
torosus, 130.
Tropidolepis scincicauda, 166.
Tropidonotus leptocephalus, 179.
ordinatus, 181.
ordinatus var. couchii, 180.
ordinatus var. hammondii, 181.
ordinatus infernalis, 180.
ordinoides, 179.
parietalis, 181.
tri-vittatus, 180.
vagrans, 181.
Turtle, Arizona Mud, 200.
California Mud, 199.
Pacific Leatherback, 198.
Pacific Mud, 199.
Water, 199.
Western Pond, 199.
Yellow Mud, 200.
U
Uma inornata, 151.
notata, 150.
rufopunctata, 151.
Uro-saurus graciosus, 156.
Uta elegans, 155.
graciosa, 156.
mearnsi, 154.
ornata, 157.
schottii, 157.
Stansburiana, 154, 156.
stansburiana elegans, 155, 156.
stansburiana hesperis, 155, 156.
stansburiana stansburiana, 154,
155, 156.
symmetrica, 157.
Uta, Graceful, 157.
Long-tailed, 156.
Ornate, 157.
Stansbury’s, 156.
Tree, 157.
Vv
Verticaria beldingi, 175.
hyperythra, 175.
hyperythra beldingi, 175.
Viperoideae, 194.
Ww
Water-dog, 130.
Water-lizard, Pacific, 130.
Whip-tail, California, 174.
Desert, 172.
Stejneger’s, 174.
Wood-frog, 145.
Worm, Blind, 170.
Worm-snake, Blue, 170.
x
Xantusia henshawi, 171.
picta, 171.
riversiana, 172.
vigilis, 171.
Xantusiidae, 171.
Xerobates agassizii, 199.
berlandieri, 200.
Xiphonura tenebrosa, 139.
Z
Zablepsis henshawi, 171.
Zamenis constrictor, 189.
constrictor flaviventris, 189.
flagelliformis frenatus, 190.
fiagellum, 190.
flagellum flagellum, 190.
grahami, 189.
lateralis, 191.
taeniatus, 192.
gy
1, Sppeac “ Octahell 19,
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UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 11, pp. 209-222, plate 13, 2 figures in text October 19, 1917
ASS DUD ORDA RACES OF THE WHITEH-
FRONTED GOOSE (ANSER ALBIFRONS)
OCCURRING IN CALIFORNIA
BY fi.
H. S. SWARTH anp HAROLD C. BRYANT
(Contribution from the Museum of Vertebrate Zoology of the University of California)
It seems almost incredible that the presence of a bird as conspic-
uous as a goose should have remained unknown to science until this
late date in a state where ornithology has been studied as intensively
as in California. But recent information proves this to have been
the ease. The fact is now established that two well-defined subspecies
of Anser albifrons occur in California during the winter months,
instead of the single race heretofore recognized.
For the material employed in demonstrating the differences be-
tween the two forms the authors are indebted primarily to Judge
F. W. Henshaw, of the Supreme Court of California, and also to
Mr. George Neale, in charge of the Sacramento office of the California
Fish and Game Commission. It was through the latter that we re-
ceived our first intimation of the existence within this state of a gray
goose different from the common species. Judge Henshaw secured
and donated to the Museum of Vertebrate Zoology a series of fourteen
specimens, sufficient in number for the prosecution of a detailed study,
and he also supplied valuable written notes upon the habits of these
two races of geese as observed in their winter home. Mr. Neale
donated two specimens, as described beyond, and also gave us written
notes regarding the habits and appearance of the birds in life.
We are under indebtedness of another sort to Dr. Barton W. Ever-
mann, director of the California Academy of Sciences, who permitted
210 University of California Publications in Zoology (Vou. 17
examination of the extensive series of white-fronted geese in the collec-
tion of that institution.
In the spring of 1916 a conversation took place between the junior
author of this paper and Mr. George Neale, in the course of which the
latter described what he called a ‘‘tule goose’’ or ‘‘timber goose,’’
distinguished from the common white-fronted goose by its much
greater size, its call notes, and certain details in its habits. In fur-
therance of our efforts to ascertain the specific identity of this large
goose we obtained during the ensuing winter, from the two donors to
whom acknowledgments are made above, specimens as listed below,
all taken in the vicinity of Butte Creek, near West Butte, Sutter
County, California.
The numerals used in the following pages for reference to speci-
mens are the collection numbers of the Museum of Vertebrate Zoology.
All measurements are in millimeters. Color terms are from Ridgeway,
1912.
From Judge Henshaw: Nine tule geese, five adult males, three
adult females, and one immature female (nos. 27175-27177, 27572,
27573, 27575-27578) ; five white-fronted geese (nos. 27574, 27579-
27582).
From Mr. Neale: Two tule geese, one entire specimen, an adult
male (no. 27134), and one specimen consisting of the head and neek
of an adult bird, preserved in alcohol (no. 27583).
There is in addition in the Museum collection, a series of thirty-six
skins of the white-fronted goose, from the vicinity of Los Banos,
Merced County, California. Examination was also made of the series
of forty-three specimens of the latter species, from the same locality,
in the collection of the California Academy of Sciences.
The first specimen received of the so-called tule goose was a male.
The great size of this individual was at first attributed to age, and
it was suggested that very old ganders of our common species might
sometimes attain exceptional dimensions, much greater than the mode.
But the acquisition of additional specimens refuted this conjecture,
for females were later secured which in spite of a notable difference
in their ages (one of the specimens being immature) were all of
approximately the same size, and much larger than the common form
of white-fronted goose.
Except for the immature plumage, which is worn for at least
the first year, the external appearance of these birds yields no reliable
clue to their age. There is a general belief, however, that the black
1917] Swarth-Bryant: White-fronted Goose in California 211
blotching of the lower parts increases in extent with the passage of
years; and the individuals occasionally encountered in which the
lower breast and abdomen are uniformly black are thought to be
of great age. If this idea is reasonable, and it appears to be so,
the big birds in question cannot be regarded as being of exceptional
age. None has the belly unusually heavily blotched, while on the
other hand several of the smaller sized birds are almost entirely
black below.
Sex and age having been thus excluded as causes of the observed
differences, it became necessary to make a careful comparison of the
two series of specimens, the large tule goose and the small white-
fronted goose, in order to find a more satisfactory solution of the
question. Fortunately, enough examples of both were available to
make this practicable.
The difference in size between the two subspecies was equally
noticeable whether the comparisons were made before skinning the
specimens or in tabulated measurements. This was at once sugges-
tive of a like contrast within the Branta canadensis group. The large
tule goose may be compared with the Canada goose, which it closely
approximates in bulk. The smaller white-fronted goose is comparable
to the Hutchins goose in size, while, to complete the analogy, the
Asiatie species, Anser erythropus, may be paralleled with the tiny
cackling goose. A similar variation is to be found in the North
American snow geese (Chen), there being in this genus three species
comparable in size.
A color distinction that is at once apparent between the two series
of Anser is that the larger birds are of a browner tint, and the smaller
ones more gray. This is especially noticeable on the heads and necks.
In some individuals of the larger race the head is extremely dark
brown, almost black. In the distinctive markings, the white face
patch and the black blotches on the belly, there appears to be no
difference between the two.
In one of his communications (see below) Judge Henshaw called
attention to the fact that the tule goose has a yellow eye-ring, a feature
that is not present in the white-fronted goose. This important char-
acter is not apparent in a dried skin, and it had been overlooked
in the first few specimens that came in, but its presence was verified
in all but one of the large geese subsequently received. This marking
is similar to that ascribed to Anser'erythropus, the edge of the eyelid
being naked skin, and forming a bright yellow ring about the eye.
212 University of California Publications in Zoology (Vou. 17
It was conspicuously present in both sexes, and even in the single
immature female; in only one specimen (no. 27575) was it absent.
In the common white-fronted goose the eyelid is dark brown.
Another character that may be noted is the number of tail
feathers. Of the six males of the larger race at hand, four have
eighteen tail feathers each, and the two which have a lesser number
appear to be molting or to have had tail feathers shot away. The
four females have each sixteen remiges. Of the series of the smaller
race, male and female alike have sixteen tail feathers. Of twenty
specimens examined none has more.
In the two lots of birds examined, we thus find differences of size,
color of plumage, number of tail feathers, and in the character of the
eye-ring. There seems to be no question but that the series are rep-
resentative of two distinct subspecies at least. In fact, to anyone
handling the birds in the flesh, the differences between the two are
obvious beyond dispute. Granting, then, the existence of two races,
the question arises as to the proper names to be applied to them.
The American white-fronted goose has long been known as Anser
albifrons gambecli Hartlaub. It would seem at first thought that the
more common North American species (and evidently one of these
two species is much more common than the other, in California at
least) should be the one to bear this name, but careful consideration
of the question makes it seem doubtful that this is the correct inter-
pretation of the facets.
Hartlaub’s description (1852, p. 7) of Anser albifrons gambeli
reads as follows:
ANSER GAMBELLI, Nob.—(Notice provisoire.)—Synon. Anser albifrons Americ.
septentr.
Nous avons examiné trois exemplaires de cette espéce d’Oie, dont deux pro-
venaient du Texas et 1’un du sud de 1’Amérique du nord. Ce dernier est presque
adulte; les deux du Texas sont des jeunes. L’énorme grosseur et la forme
différente du bee nous force de séparer cette espéce de notre albifrons. Voici les
dimensions comparatives:
A. GAMBELLI A. ALBIFRONS
TWomeaibes POStri cay che peree te senaceteee eee 2s GAL Oar
TACIT Cb: eeseteueesusestesee av 4r"u LS Bi
Altitud. rostri later i ile rte OMA,
Circumferent. rostri (ad! bas? cesses Gi PH eae A
Longit. tars. PET eyo ee Ply ESB,
Diowmedye PA (en Oa
Le congrés des ornithologistes, 4 Berlin, en 1851, a approuvé la séparation
spécifique de cette Oie américaine.
1917 | Swarth-Bryant: White-fronted Goose in California 213
As regards most of the measurements given in the above descrip-
tion, it is apparently impossible to ascertain the exact manner in
which they were taken. Hence it is difficult to use them in making
comparisons. One of Hartlaub’s measurements, (‘‘Longit. rostri a
Fig. A. Anser albifrons albifrons, adult male, no. 21922, Los Banos, Merced
County, California; natural size.
Fig. B. Anser albifrons gambeli, adult male, no. 27573, West Butte, Sutter
County, California; natural size.
fr.’’?) may safely be assumed as corresponding to length of culmen
as we have measured it. This measurement, as given for his A.
Gambelli, translated into millimeters (58 mm.), is within the range
of variation of our larger goose. It can not be applied to the bill of
the smaller variety.
214 University of California Publications in Zoology | Vou. 17
Howsoever they were taken, Hartlaub’s measurements show about
the same proportional differences between the two races he had in
hand, as there are in our two sets of birds. His new species was
evidently a very large bird, and one, at least, of the measurements
he gives of it can be applied to the larger of our two forms. It seems
safe to say, therefore, that the name Anser albifrons gambeli should
be used for our large tule goose.
The question then arises as to the status of our small white-fronted
goose, which is evidently by far the more common of the two in
California. European writers have been reluctant to recognize an
American race of Anser albifrons. Salvadori (1895, p. 97) describes
gambeli as ‘‘searecely different from A. albifrons; on the average it
is larger and has a bigger bill.’’ Alphéraky (1905, p. 42) refuses to
recognize gambeli at all, though conceding that occasional specimens
from North America attain a greater size than any European birds.
Of deseriptions and measurements as given in most of the American
literature on the subject, it may be said that the diagnoses are not
convincing as proofs of the subspecifie identity of Anser a. gambeli,
though there is throughout recurrent mention of occasional unusually
large sized birds. (In this connection see Coues, 1874, p. 547.) It
seems likely that confusion has arisen through failure to discriminate
between two perfectly distinct races, and that the explanation of the
puzzle is as follows: That gambeli exists as a distinguishable North
American subspecies of large size, as originally described, but that
there also exists in North America another form of smaller size, and
that the two occur together during the winter months. The smaller
bird is, to all appearances, indistinguishable from the European form.
In other words, the race of the white-fronted goose which is most
common on the Pacific coast of North America is Anser albifrons albi-
frons (Seopoli).
In testing this theory reference should be made to the aecompany-
ing tables. The measurements of European A. albifrons albifrons
are taken from Alphéraky’s (1905, p. 46) careful study of the
species. The California series of this subspecies used in comparison
was collected at Los Banos, Merced County, during the winter of
1911-12. It will be noted that the measurements of this last series
fall within the extremes given for the European birds. No European
or Asiatic specimens are available for actual comparison, but no differ-
ences, save of size, are claimed to exist between these and American
birds, and as it is evident that there are no size differences between
1917] Swarth-Bryant: White-fronted Goose in California 215
the series here compared, it seems justifiable to apply to the smaller
American race the name of the common European subspecies, Anser
albifrons albifrons.
The differences existing between the two subspecies may be sum-
marized as follows:
Anser albifrons albifrons
Size small (wing 384-422); bill small (culmen 44-52); tail feathers, six-
teen; coloration in general paler, head and neck grayish; naked skin
at edge of eyelid, grayish brown.
Anser albifrons gambeli
Size large (wing 420-475); bill large (eulmen 53-62) ; coloration in gen-
eral darker, neck dark brown, head blackish; tail feathers, male eighteen,
female sixteen; naked skin at edge of eyelid, yellow or orange.
In colors of ‘‘soft parts,’’ before the birds are skinned, there were
no distinguishable differences between the two series in eyes, bill or
feet. The eyelids of the two varied as already pointed out. Accord-
ing to Stejneger (1885, p. 146) appreciable changes occur within an
hour after death, and as probably none of our specimens came to hand
until after a lapse of at least twenty-four hours, the colors as we noted
them, particularly of the bill, may be quite different from those of
the living bird. There were great changes, however, in these parts
after the prepared skins had begun to dry. “Some faded and some
darkened.
An adult male of Anser a. gambeli (no. 27134) was colored as
follows: Upper mandible, general ground-color, light buff tinged with
purplish along culmen and at the edges; lower mandible, upper edge
of rami, purplish, lower edge, yellowish; naked skin between rami,
light buff; feet (tarsus, toes and web), ochraceous salmon. The color
of eyes in all the specimens was dark brown.
As of general interest it is worth while to record here the fact that
all the birds taken in January were molting extensively, over head,
neck and body. There was evidently a general freshening of plum-
age, apparently involving everything but flight feathers to a greater
or less extent. Newly appearing black feathers on the lower parts
were particularly noticeable.
The present study is based entirely upon specimens collected in
winter. Not a single breeding bird is available for comparison, un-
fortunately, so that we are unable to indicate the summer ranges of
the two North American forms.
The following theoretical breeding ranges are suggested by the
facts thus far ascertained, though demonstration of the truth of the
216 University of California Publications in Zoology [| Vou. 17
hypothesis must depend upon future investigation. With Anser albi-
frons albifrons occurring commonly in the western United States
during the winter months, it would seem fair to assume that the
known summer range of the subspecies, covering northern Europe
and Asia, also extends continuously over western Alaska, and for an
undetermined distance eastward. Pursuing this hypothesis, and
assuming, as we have the right to do, that the two races do not occur
together during the breeding season, we may infer that the summer
home of the large A. a. gambeli is restricted to points farther eastward
in Arctic America than the region inhabited by A. a. albifrons. This
distribution would explain the relative scarcity of the first mentioned
upon the Pacifie Coast. Study of specimens from the Mississippi
Valley.and points farther east should go far toward confirming or
refuting this theory, for upon the hypothesis advanced, the condition
existing there, must be the opposite of that obtaining in California.
There should be, namely, an abundance of the larger Anser a. gambeli,
and a searcity of the smaller race. In this connection it is of interest
to note a comment made by Nelson (1877, p. 186) upon specimens
from Illinois: ‘‘I have examined a number of specimens, which by
correct comparison were at least one-fourth smaller than the average.”’
The white-fronted goose is known, of course, to breed commonly
in Alaska, but there are no Alaskan skins at hand for comparison;
nor have we been able to discover published measurements of speci-—
mens either from this section or elsewhere in America, in which
dimensions of specified individuals are given together with explicit
statements of exact place of capture. Consequently the assumption
that the breeding bird of western Alaska is identical with the smaller
of the two subspecies visiting California in winter, is an unproven
hypothesis. It ean be said, however, that measurements of eggs from
the Yukon region, as given by Nelson (1887, p. 83) agree reasonably
well with the dimensions given by Alphéraky (1905, p. 56) for those
of the European bird. Eggs of a set in the Museum of Vertebrate
Zoology (no. 714) from Cape Vancouver, Alaska, are close to the
minimum dimensions given by Alphéraky. It would seem that eggs
of the large sized A. gambeli should be measurably larger than those
of A. albifrons.
In considering the possible continuity of range of Anser a. albi-
frons over Asia and Alaska, mention must be made of a goose taken
by Stejneger (1885, p. 145) upon Bering Island, which he records
as Anser a. gambeli, saying that ‘‘it matches average North American
1917] Swarth-Bryant: White-fronted Goose in California 217
oy
specimens in every particular. The following details of this bird
can be used in comparison with our data: Sex, female; total length,
685 mm.; wing, 417; tail feathers, 124; bill, from tip to frontal
feathering, 51; tarsus, 78; middle toe with claw, 76; weight, 634
pounds (fat); naked eye-ring, dark brownish gray.
It is, of course, uncertain whether these measurements were taken
in the same manner.as our own, but, disregarding this possibility, it
will be seen from the figures given that the total leneth accords with
that noted by us for the smaller American race, which we eall albifrons,
while the others are all intermediate between the two. Stejneger,
with his customary painstaking accuracy, earefully records the color
of the naked eye-ring, most fortunately so, as it appears to be a valu-
able character. The fact that it is dark brownish gray in the
specimen in question seems, with little doubt, to stamp the bird as
Anser a. albifrons.
On the whole, while concurring with this author that his Bering
Island white-fronted goose agrees with average North American birds
in its characteristics, we believe it belongs to the smaller, apparently
the more common, of the two American races. Its slightly greater
size, as compared with most European A. albifrons, is in accord with
Alphéraky’s finding of an inerease in the size of birds from eastern
Asia, as compared with European specimens.
All of the examples of tule geese at hand came from a limited
region in the Sacramento Valley, in the vicinity of Butte Creek and
Butte Slough, in Sutter County. While the bird is apparently of
fair abundance in this region in winter, we have little data demen-
strating its presence at any other point in the state. In the extensive
series of A. albifrons albifrons in the collections of the Museum of
Vertebrate Zoology and of the California Academy of Sciences, all
taken in the vicinity of Los Banos, Mereed County, in the San Joaquin
Valley, there is not a single example of the larger bird. There is,
however, a persistent rumor among market-hunters of the Los Banos
district to the effect that a large form of white-fronted goose exists
and has been killed there. In a letter received from Mr. George
Neale, the statement is made that Mr. A. W. Stuart, of Grand Island,
once killed two large gray geese, ‘‘as large as honkers,’’ at Maine
Prairie, Solano County.
In the Sacramento Valley, market-hunters and the sportsmen of
the gun clubs alike affirm the existence of two races of the white-
fronted goose, differing in appearance, habits and call notes. It is
218 University of California Publications in Zoology [V0. i
said that the two kinds flock separately, for the most part; and that
the larger race is never seen in such big flocks as is customary with
the other, but is most frequently noted singly or in pairs. Also that
while the smaller variety is a common frequenter of grain fields and
uplands generally, the larger one is pre-eminently a denizen of open
water or of ponds and sloughs surrounded by tules and willows. The
predilection of the latter species for such localities has given rise to
the local names by which it is known, ‘‘tule goose’’ or ‘‘timber goose,”’
as contrasted with the upland-frequenting ‘‘speckle-belly.’’
The habits and appearance in life, of the tule goose are described
in the following excerpts from a letter written by Judge F..W. Hen-
shaw to Dr. J. Grinnell, director of the Museum of Vertebrate
Zoology :
On Monday last, Jan. 22nd, accompanied by my friend Sam Lamme, I went
out to secure for you some tule geese on the grounds of the West Butte Country
Club in the Sutter Basin. We went by boat into the more unfrequented and
inaccéssible parts of this lake and there shot for you four tule geese and five
American white-fronted geese. Sam Lamme is remarkable even for a professional
hunter. His ability to call wild fowls of all kinds is little short of marvelous,
and we could easily have killed more of these birds, but we stopped when we
had secured the number that you desired. We were out only during the morning,
and in that time (accepting Sam’s verdict as to their character) we certainly
saw over 150 tule geese. Usually they were single birds or in pairs, though at
times we would see flocks of eight, ten, twelve, or sixteen. At times also we saw
mixed flocks. Sam explained this by saying that the tule geese never joined the
smaller white-fronted geese, but that the latter would frequently attach themselves
to a pair or to a flock of tule geese and trail on behind. I was myself a witness
to the joining of these birds upon several occasions. The tule geese were always
in the lead, paying no attention to the other geese which joined them, and in
turn the other geese would frequently leave the tule geese after accompanying
them for a short time. When the tule geese were by themselves and at a distance
it was difficult for me to tell with any certainty whether they were tule geese or
the ordinary white-fronted geese, but when the two kinds were together the differ-
ence was most plain and showed not alone in size but in the conspicuously longer
neck of the tule geese. Also the notes of the tule goose, according to Sam, while
similar, are of different quality from those of the white-fronted goose, and while
I was unable to detect the difference myself, he frequently verified his own nicer
sense of hearing. The difference he described by saying that the notes of the
tule goose were coarser and harsher. His nicety of ear I had him demonstrate
upon several occasions. He would say upon hearing the call of a bird, ‘‘that is
a tule goose,’’ or ‘‘that is a gray goose,’’ and I would reply, ‘‘call him in, Sam,
and let’s make sure.’? When he did so, in every instance his judgment proved
to be correct. In explanation of this, let me say that he called many of both
varieties within range of our guns after we had killed all that you required, so
that in many instances I had to base my conclusion upon observation of the birds
in the air. Another noticeable fact was that the tule geese, while shy in the
sense that they resorted to the more remote parts of the marsh—the white-fronted
1917] Swarth-Bryant: White-fronted Goose in California 219
geese being everywhere by the hundreds—were much more confiding and answered
much more readily to Sam’s call, coming in directly and without the usual wary
circling. Several times, for example, it happened that in a mixed flock the
white-fronted geese would turn and leave, while the tule geese would come sailing
on to what would haye been their destruction.
I mentioned to you over the phone the conspicuous bright orange membrane
fully surrounding the eye of the tule goose. It did not appear upon any specimen
of the white-fronted goose, and Sam declared that he had never seen it on a
white-fronted goose. Sam, I should add, was born in Sutter county, has been for
years a market hunter, and is exceptionally endowed with powers of observation
and wild fowl mimicry.
Transmitted May 1, 1917.
220
University of California Publications in Zoology
[ Vou. 17
MEASUREMENTS IN MILLIMETERS (AVERAGE, MINIMUM, AND MAXIMUM) OF THE
Races or Anser albifrons
Wing
(430.0-475.0)
6 male Anser albifrons
gambeli; California..
10 male Anser albifrons
albifrons ; California
4 female Anser albifrons
447.2
409.6 (384.0-422.0)
gambeli; Calitornia.. 430.5 (420.0-440.0)
10 female Anser albi-
frons albifrons; Cal-
THROW ANE, © ceeredeeesercare ice 394.5 (384.0-404.0)
Adult Anser albifrons al-
bifrons; Old World 375.0—435.0"
Height of bill
6 male Anser albifrons gam-
beli; California ...
10 male Anser albifrons albi-
mows Oplbhdaie, 23.3 (21.5-26.0)
4 female Anser albifrons gam-
beli; California
10 female Anser albifrons al-
bifrons ; California ............
Adult Anser albifrons albi-
rons «Olde nvorl disse 23.57
26.5 (25.0-28.0)
25.7 (25.0-27.0)
20.4 (18.0-21.5)
Total length?
Cali-
6 male Anser albifrons gambeli;
fornia _ ..-- 830.5 (810.0-854.0)
2 male Anser albifrons albifrons; Cali-
OLN sare sees eee ess ook eee 733.0 (730.0-736.0)
4 female Anser albifrons gambeli; Cali-
POPMIG, H Se sis peepee tie eta reeseemetee seer 779.7 (745.0-797.0)
3 female Anser albifrons albifrons;
Galli orm ae ceeee stone ce eeee reer 687.6 (685.0-692.0)
Adult Anser albifrons albifrons; Old
Word dy Sees ocec ccs teccsctsceseaeeenaee sae nner 635.0—-760.0"
Tail
135.5 (124.0-144.0)
121.2 (107.0-135.0)
127.2 (110.0-135.0)
119.6 (113.0-126.0)
Tarsus
81.9 (80.0-84.0)
73.6 (71.0-79.0)
79.0 (77.0-83.0)
70.1 (64.0-73.0)
51.0-81.0*
Culmen
60.1 (57.0-62.0)
49.6 (46.5-52.0)
55.6 (53.0-58.0)
46.0 (44,0-48.0)
40.0-56.0
Middle toe without
claw
79.6 (73.0-84.0)
67.1 (61.0-73.0)
75.0 (73.0-80.0)
63.1 (58.0-68.0)
Spread wings?
1623.6 (1560.0-1670.0)
1476.5 (1471.0-1482.0)
1572.0 (1510.0-1661.0)
1405.3 (1384.0-1437.0)
WEIGHTS (AVERAGE, MINIMUM, AND MAXIMUM) OF THE RACES OF Anser albifrons
6 male Anser albifrons gambeli; California -.....
2 male Anser albifrons albifrons; California ....
4 female Anser albifrons gambeli; California -...
3 female Anser albifrons albifrons; California
Adult Anser albifrons albifrons; Old World -...
6 Ibs.
4 Ibs.
1 Alphéraky, 1905, p. 46.
* Measured prior to skinning.
* Extremes of twenty-one specimens; Alphéraky, loc. cit.
7 Ibs. 4 oz. (7 lbs. 1 02-7 Ibs. 8 oz.)
5 Ibs. 4 oz. (5 Ibs.-5 Ibs. 6 oz.)
5 oz. (5 lbs. 5 02-7 Ibs.)
12 oz. (3 Ibs. 14 0z.-5 lbs. 8 02.)
4 lbs.—6 lbs.*
1917] Swarth-Bryant: White-fronted Goose in California 221
LITERATURE CITED
ALPHERAKY, 8.
1905. The geese of Europe and Asia. London, Ward, vi+195, frontispiece,
24 pls.
Cougs, E.
1874. Birds of the Northwest: a hand-book of the ornithology of the region
drained by the Missouri River and its tributaries. U.S. Geol. Sury.
Terr., Mise. Publ., 3, xi+ 791.
HArTLAus, G.
1852. Descriptions de quelques nouvelles espéces d’Oiseaux. Paris, Rev. Mag.
-Zool. (2), 4, 3-9.
NE.LSon, E. W.
1877. Birds of northeastern Illinois. Bull. Essex Inst., 8, 90-155.
1887. Report upon natural history collections made in Alaska between the
years 1877 and 1881. Arctic Series of Publications issued in con-
nection with the Signal Service, U. S. Army, 3, 337, 21 pls.
Ripeway, R.
1912. Color standards and color nomenclature. Washington, D. C., Ridgway,
iii + 44, 53 pls.
SALVApDoRI, T.
1895. Catalogue of the birds in the British Museum, Chenomorphae (etc.),
27, xv + 636, 19 pls.
STEJNEGER, L.
1885. Results of ornithological explorations in the Commander Islands and
in Kamtschatka. U.S. Nat. Mus., Bull. 29, 382, 8 pls., several figs.
in text.
PLATE 13
Skins of adult males of Anser albifrons albifrons (at left), Mus. Vert. Zool.,
no. 27581, and Anser albifrons vane cn right), Mus. Vert. Zool., no. 27134;
photographed on same scale.
These specimens are fairly representative of the size differences existing
between the two forms. Both were prepared by the same person, and care was
exercised that neither should be unduly lengthened or shortened. In each case
the dried study skin measures in total length within a few millimeters of the same
measurement as taken before the bird was skinned.
[222]
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UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 12, pp. 223-404, plates 14-24, 24 figuresintext January 31, 1918
A SYNOPSIS OF THE BATS OF CALIFORNIA
BY
HILDA WOOD GRINNELL
(Contribution from the Museum of Vertebrate Zoology of the University of California)
CONTENTS
PAGE
Tim GTCTOR © tense inn Be ee nee ee pe RU ee pS Saar ee SEE SAR RR re eee 225
BENG Feo 0 Wy) © Cl 70 ts sce 225
NEE ee a er eee ao 226
INGESTED TOTTI eo cn eee ce er ee eee te re a eR RS er ee Reiter ter ora 227
ANG cl a0 seg Es Se eval rE aS ees ee a rece ee ee ese aera 228
(Seg aVS SREY uit ed AAAS em Se he PE omc 228
TORY CSHIEUNG | Gece cceceei cose tipi ae eee eee eee Eo BERR eee ener Rea eee 229
TEIGEATRIYED see ceneseretREe oRES eeESSL PEE RES REE Ec ESS RPE 229
ae ee a ee a Nog ea ce dat aes oer Sepe seee sae neat sae Reman erence cadssesnenaenan 230
aes 231
231
TRUS] aS) One. UE HST ee ae ea Eee 232
ICV ETON, cess ee eee ee a OE ESP CO Er ee . 232
Tfoyayal carol) Theravada ess 1G) ON NS) ee ee rarer ER ee SEE ec aEE RoE . 235
Tearearalriayss TBI OEY seekers eat ee ec een rer oer Ecc coe Sere cor eS ee eee Cee . 236
NWO esse ee Se IE ee ee ee . 206
TD paveyannGyss OME VES) eer eee eso or 237
Economie Value of Bats and their Conservation —...........-..------------------------eeee 238
C0) aay erat ek © SUE TATA CA Speen eee ener ee ce eran 240
(Champs O MG IDS ret OHTA CONE BEE Se ee 241
TOYS occsee eae se oes Se ne SSS EPS a oR ae RCE EEE 243,
(CHAOS ENCOUN, + cee eee eee ee aro SS a rr eee 245
ENG NYEERETEER OY, cs eee Re EPP 244
PS essrctr a Op Wena ear Oa een etna esas aun ogee natn et ea nwa ae ema ee Re rc apse eee eat naee cen 244
Nomenclature : .. 245
Classification yt: eset =: 245
Keys etor identification of (Calitormiay (eats) 2e rece. necnsscecteeaecareesamcevenece wwe nerenesecreearen= 246
Table showing Comparative Sizes of California Bats -.........--------------------- 251
@rdembs Chino p tier ay geese sascese ones ees n eee anne Sees ov sees ann sctees seer enscsveseeseeanesetncesteagen-sooeunsecerteus 252
MUD OLdereMacrOChiNG Pp LCL eccens ceca cee ease ceca esac aca oe tennn comes eocewsocenn dec nente ness toarecrezunere 252
224 University of California Publications in Zoology [Vou.17
amily MeL yO stomiid aap jens acescecseece testes nme ene ee esa ee eee 252
Genus Macrotus ..............-
Macrotus californicus
Family Vespertilionidae -..
Subfamily Vespertilioninae -
Genus Myotis ..... 258
Myotis velifer ... - 259
Myotis occultus -.............--- - 261
Myotis lucifugus altipetens ..
Myotis lucifugus alascensis ..
Myotis longicrus longicrus .
Myotis longicrus interior .... 271
Myotis yumanensis yumanensis 273
Myotis yumanensis sociabilis -. 276
Myotis yumanensis saturatus ... 278
Myotis californicus californicus -. 279
Myotis californicus quercinus ... 285
Myotis californicus pallidus .. =F 1288
Myotis: OVtmOMUs: q2.<. ssn - ccc wach ans ose eos 290
Myotis evotis -..... oe ho ee 291
DMYOUS® THYSGNOGES asco nica sccckescct acco ees 297
Genus Dasionycteris 22-2 22. q.cssecan nto ecco se 299
LGStonycterts: MOCCWWAGGNS <2. ee ee 300
Genns ePipistrellush seen ee 304
Pipistretius hesperus) hesp ers oon... ose ee 305
Papistretlus Wesperus MenTVamne sxe onsen 311
Genus: Hptesicus) 2cc.s.0s0 css ee ee 314
Eptesicus fuscus ...
Genus: Nyctenis -.2i..c5.0.8 oo eae eee
Nycteris: borealis’ teliotts..=... oe 323
Naycterts CNET CG, <2....08. Soc tenet ee ee 330
Genus Buderma: «2.25... 5 nee te ee ee 336
Huderma smaculatum) «025... SS eee 336
Genus Corynorhinus | 5 2 ee 339
Corynorhinus vafinesqua paliescens ee eee 340
Corynorhinus rafinesquit intermedius -...-.-----.---c0--eeeeeeeeeeeeeceeceeeeee eee 344
Subfamily -Nyctophilinae, =: — s.r 347
Genus’ -Amtrozous): <= 2c ee 347
Antrosous! PAQlUNdus sscci cco 348
ANtTOBOUS! DGOCVCUS oe cc Fock eee 352
Family ‘Molossi@ae \n028 22. scc sooo ee 357
Genus) INycbimomi us ace 2 acc Sc ca cece tek ee ee 357
INGetInOMUs <depressus .x<- ccs scccc sce ss eo 357
Nayetinomus: fEMOTOSACCUS 28. - fcc cose 360
Nyctinomus) mexicanus = <.<0.2- 0 ee 361
Gettus: umops) +225 See 369
Humops caltfornicus 2=-.5i< 23) Se ee 370
Diterature iCited: 1.5.2... ae 376
1918 | Grinnell: A Synopsis of the Bats of California 225
INTRODUCTION
The paper here presented is the result of the writer’s endeavor
to gather together, and to add to, the facts already known concerning
the distribution and habits of the California representatives of one
of the most interesting, though least known, groups of mammals,
the bats. The records of palaeontology show the great antiquity of
the order to which the bat belongs; indeed, the bat had become master
of the air long before man walked upright. The first fire the cave man
lighted in his rocky refuge revealed to him the ‘‘little upside-down
bat’’ clinging to the roof of his cavern, and doubtless he knew more
of its habits than we do today. In the course of ages, as man with-
drew himself more and more from contact with the wild creatures
and became more imaginative, he grew to fear this haunter of caves
and dweller in darkness. Shakespeare but voiced the popular senti-
ment of his day when he elassed the bat with toads, snakes, and newts,
a fit ingredient for the witches’ brew. In reality, however, the bat is
a warm-blooded, friendly little creature, as deserving of our good will
as the bird whose place he takes at twilight, when he skims the air
for those insects which escape the vigilance of the bird by reason of
their time of flight.
The number of students of natural history within our state is
rapidly increasing and it is hoped that by turning their attention
more generally toward our bats, this paper may be the means of
extending our knowledge of these animals far beyond the limits of
the present summary. The facts gathered together here have been
derived from the following sources: All the available published liter-
ature; the data attached to specimens ; the many volumes of collectors’
manuscript field notes on file in the California Museum of Vertebrate
Zoology; and the writer’s own study of specimens in the Museum
and of live bats in the field. As a rule, the source of facts cited on
authority is designated. All specimens listed are contained in the
Museum of Vertebrate Zoology, except where their location is other-
wise indicated.
ACKNOWLEDGMENTS
For permission to examine the collections under their charge and
to make use of data derived from them, the writer is indebted to
Dr. Charles H. Gilbert and Professor John O. Snyder of Leland Stan-
226 University of California Publications in Zoology [Vou.17
ford Junior University; to Dr. Barton W. Evermann and Mr. John
Rowley of the California Academy of Sciences; to Professor Charles
A. Kofoid of the University of California; to Dr. Leonard Stejneger,
Mr. Gerrit S. Miller, Jr., and Mr. Ned Hollister of the United States
National Museum; to Mr. Henry W. Henshaw, Mr. E. W. Nelson and
Mr. Arthur H. Howell of the United States Bureau of Biological
Survey; to Dr. Witmer Stone of the Academy of Natural Sciences
of Philadelphia; to Dr. J. A. Allen of the American Museum of
Natural History, New York; to Dr. Glover M. Allen of the Museum
of Comparative Zoology, Cambridge; and to Mr. Perey A. Taverner,
of the Museum of the Canadian Geological Survey, Ottawa. To Dr.
Joseph Grinnell of the California Museum of Vertebrate Zoology
the writer is indebted not only for permission to make full use of
the collections and field notes under his charge, but also for encourage-
ment and frequent help given during the five years in which this
paper has been in preparation.
A preliminary manuscript, the first draft of the present paper,
was prepared when the writer was a graduate student in the Depart-
ment of Zoology of the University of California, m 1912-13; during
that year much kindly criticism was received from Professor Charles
A. Kofoid, head of the department.
To Mr. Charles L. Camp, Mr. Wilson C. Hanna, Miss Grace Swerd-
feger, Miss Winifred N. Wear, and Messrs. Eldon, Adrey, and Raleigh
Borell, the writer is under obligation for contributions of living speci-
mens of bats.
Mr. Frank Stephens has generously permitted the use of notes
gathered during many years of field study of California mammals.
Finally, the author is deeply indebted to the entire staff of the
California Museum of Vertebrate Zoology for aid given in many ways.
The facilities afforded in this institution for this special study have
been indispensable.
MaTERIAL
The Californian material studied, and upon which this paper is
based, consists of some 1500 specimens, from the following sources:
942 specimens from the Museum of Vertebrate Zoology; 218 from
the collections of Leland Stanford Junior University; 107 from the
United States National Museum; 88 from the collection of the United
States Bureau of Biological Survey; 34 from the collection of the
1918] Grinnell: A Synopsis of the Bats of Califorma 227
Academy of Natural Sciences, Philadelphia; 20 from the collection
of the Department of Zoology of the University of California; 20
from the American Museum of Natural History, New York; 17 from
the collection of the California Academy of Sciences, San Francisco ;
9 from the Museum of Comparative Zoology, Cambridge; 9 from the
collection of the San Diego Society of Natural History; 5 from the
collection of Frank Stephens; and 1 specimen from the collection of
Rea Weakoch
Many specimens of bats from outside of the State of California
have been used in comparison, and these are also from the above-
named sources, and from the Museum of the Canadian Geological
Survey, Ottawa.
The material examined has consisted chiefly of dry skins with
skulls; but it has generally been possible to secure fresh and alcoholic
specimens from which to derive the essential supplementary data.
MEASUREMENTS
Throughout this paper all measurements of total length, tail
vertebrae, and foot are as taken in the flesh by the collector, unless
otherwise stated. These are likely to be somewhat longer than meas-
urements obtainable from dried skins or from alcoholic specimens.
Since averages are useful only for comparison with other averages,
detailed measurements of specimens are given wherever possible. In
selecting specimens for measurement examples have been chosen which
show extremes of variation, as far as contained in the material at hand.
All measurements are stated in millimeters. In the fresh specimen
it 1s customary to measure the total length, and lengths of tail, foot,
ear, and tragus. To do this with facility, the bat should first be laid
out flat upon its back, when the total length from end of nose to tip
of tail, may be taken. When the tail is to be measured one point of
a pair of dividers should be set on the rump at the extreme base of
the tail and the other point placed at the tip of the straightened tail.
The foot should be measured from the tip of the longest claw to the
upper edge of the heel. Two methods of measuring the ear are used
and as these differ somewhat in results the particular one used should
always be specified. One is ‘‘ear from crown,”’ and is taken with
one point of the dividers set on the skull on the inner (convex) side
of the ear and the other at the tip of the ear. The second method,
and one which results in a slightly longer measurement, is ‘‘ear from
228 University of California Publications in Zoology [Vou.17
meatus,’’ one point of the dividers being placed in the meatus, the
other at the tip of the skin of the ear, as before. The first-named
method of measuring the ear is recommended as it seems to admit
less chance of error. By either method the measurement is, of course,
of more value when taken from the fresh (unskinned) animal.
Mopes oF PRESERVING Bats
Two modes of preserving bats as specimens are used by collectors,
namely, immersion in an aleohol or formalin solution of sufficient
strength to arrest decay and yet not of such concentration as to harden
the tissues unduly; and preservation as dry stuffed skins, with the
skull cleaned and saved separately. The latter method is that usually
employed in preserving other mammals.
Since the color of the fur of bats is of great importance in
specific and subspecifie determinations, and since the skulls are of
prime importance in the matter of general identification, the second
method is to be recommended in most cases. <A bat skin should be
made up in the same way as are the skins of other small mammals,
save that no wire should be used in the wings or feet, the tail only
being wired. The wine bone should be cut at about the middle of
the humerus and the distal portion of that bone, together with all the
other wing bones, left in position. The femur should be cut at the
middle and its distal portion, together with all bones of the lower leg
(tibia) and foot, left in position. In ‘‘setting’’ a bat skin, the wings
should be neatly folded along the sides and held so by pins at the
sides (but not through the membranes) until dry, the interfemoral
membrane should be well spread, the feet turned shghtly outward,
and the knee joint slightly bent. Observance of these precautions will
facilitate the measuring of bone lengths in the dry skin.
Where several specimens of the same species of bat are secured,
one or more may be preserved as alcoholics, since the form of the
wings and ears are more easily studied in bats preserved in this way.
SENSES OF Bats
The writer of the present paper has had no opportunity to carry
on experiments to determine the relative value of the different
sensory adaptions of bats; so there are here quoted the conclusions of
Hahn (1908, pp. 185-193) and of Ackert (1914, pp. 301-343).
1918 | Grinnell: A Synopsis of the Bats of California 229
Eyesight —Hahn (loc. cit., p. 155) says of the sight of bats:
After extended observations on the subject, [ am still unable to form any
definite conclusions with regard to the importance of sight to these animals.
That they can see light and darkness and moving objects is unquestionable. That
the sense of sight is not highly developed is equally certain. The behavior of
some of the animals appears to indicate that at times they depend on this sense
to a considerable degree, both in securing food and in avoiding objects.
J. Grinnell (1913a, pp. 344-345) suggests that the fact that a
given species of bat will appear abroad at a certain time almost to
the minute each evening shows that these animals can appreciate light
intensity within a very narrow range.
It is suggested by Hahn (1908, p. 157) that a bat dipping to
drink water from a quict pool is probably attracted to it by sight, but
that the rising of moisture-laden air also helps the animal to locate
water.
Hearing —Low-pitehed rumbling noises have no effect upon bats;
but to vibrations of high frequeney these animals are extremely
sensitive (Hahn, loc. cit., p. 155). This is as one would expect to find
it, since the hum produeed by flying insects is usually high-pitched,
and the voices of the bats themselves are very shrill. .
Hahn (loc. cit., p. 156) found that as a rule the bats he had under
observation paid no attention to worms held near them, as long as
the worms remained quiet, but became excited as soon as the worms
began to wriggle. As this occurred when the bats were not touched
by the worms and when they were out of the range of vision, Hahn
infers that the food must have been perceived through the tactile
organs, these being stimulated by air currents set in motion by the
moving worms. In recording his experiments upon captive bats Hahn
(loc. cit., p. 178) says: ‘‘Sound associations are formed readily. A
sucking noise made by the lips at first alarmed the animals, but they
soon learned to associate it with feeding. On hearing it they would
look about and snap at any object that could be mistaken for food.’’
J. Grinnell (1913a, pp. 344-345) suggests that bats hunt their
insect food by sound. He says: ‘‘Even the wing-strokes of a tiny
miller must be distinetly audible to the bat which snaps it up so
unerringly. And the droning of a June beetle must sound to the bat
as penetrating as the roar of a biplane motor does to us.”’
At least one species of bat, Antrozous pacificus, undoubtedly
secures a portion of its food upon the earth, since it brings to the
roost wingless Jerusalem crickets (Stenopelmatus). It seems prob-
230 University of California Publications in Zoology [Vou.17
able that the bat is attracted to these insects by the noise which they
make in crawling about over the ground or, less probably, by air
currents due directly to the movements of the insects.
Touch.—For more than a century naturalists have been aware of
the dexterity with which blinded insectivorous bats when in flight
avoid obstacles, and the suggestion has been repeatedly made that
* which makes them cog-
nizant of the adjacency of objects which they neither see, hear, nor
ce
these little animals possess a ‘‘sixth sense,’
touch. Many experiments have been carried on in the hope of
locating this sense. Hahn (1908, p. 191) states that obstacles are
perceived chiefly through sense organs located in the internal ear,
basing his belief upon the results of experiments in which the external
auditory meatus of each bat used was filled with hardened plaster of
paris. Ackert (1914, p. 329) regards the experiments of Hahn as
inconclusive, believing that a bat so mutilated might not act in a
normal manner.
Ackert (loc. cit.) himself publishes the results of a search made
for sensory structures in the skin of bats. The species used by Ackert
were Myotis lucifugus and Myotis subulatus, species not occurring
in California, but members of our most abundantly represented genus.
He performed no experiments upon lying material, but confined his
investigations to the study of prepared sections of the tissues. <A
review of the literature upon the subject inclined him to the view
that condensations (pressures) of the atmosphere set up between an
obstacle and a bat stimulate sensory structures in the integument of
the bat. These structures Ackert suggests would have to meet the
two conditions of distribution over the parts of the bat foremost im
flight, and of superficial location, as stimulations from air pressures
are doubtless very slight.
Two types of sensory end organs found by Ackert (1914, pp. 330-
331) im the skin of bats seem to him to meet the requirements men-
tioned. The first of these consists of free nerve terminations (end-
knobs) found in enormous numbers near the surface of the epidermis.
Second, are the superficial nerve rings (and their terminal fibers),
which are so situated about the necks of the hair follicles as to be
affected by even the slightest movement of the hairs. This investiga-
tor states that the area of the integument supplied by superficial
nerve rings is insignificant in comparison with the area supplied with
nerve end-knobs. Likewise, the number of terminal fibers of the
rings is not to be compared with the enormous number of end-knobs
in the epidermis. He remarks, finally, that
1918] Grinnell: A Synopsis of the Bats of California 231
Tt is not, of course, to be inferred that all the free end-knobs function alone
as pressure perceptors, for, as is well known, the sensory nerves of the human skin
mediate at least four different qualities of sensations, namely, pressure, warmth,
cold, and pain. But the number of nerve end-knobs in the skin is so great, and
the latter in the bat is so sensitive to delicate tactile stimuli, that the number
of free nerve terminations in the epidermis functioning as pressure perceptors
must necessarily be very large.
I have repeatedly watched captive bats, of several species, when
loosed in a many-windowed room, and though they were clearly seek-
ing a means of escape, not one was ever seen to dash against a window
pane, as does a bird under like circumstances. These bats were in
full possession of the faculties of sight and hearing. To judge from
the actions of birds similarly situated, neither faculty could apprise
these creatures of the fact that glass is an impenetrable barrier. The
only theory which seems adequately to explain the bats’ recognition
of the invisible barrier, glass, is that condensations (pressures) of
the atmosphere set up between the window-pane and the bats stimu-
late sensory structures in the integument of the bats, as suggested by
Ackert, or possibly, in the internal ear, as Hahn considered probable.
According to Barrett-Hamilton (1911, p. 42), we must give Cuvier
eredit for this theory. He says: ‘‘It seems that Cuvier was not far
wrong when he wrote that ‘it is by means of the pulsations of the
wines on the air that the propinquity of solid bodies is perceived, by
the manner in which air reacts upon their surface.’ ”’
Smell—Hahn (1908, p. 154) suggests that since bats catch their
food in the air where a flying insect leaves no permanent path and
cannot be definitely localized by its odor we must infer that the bat
in seeking food does not rely on its sense of smell. But he adds: ‘‘It
must not be inferred... that the sense of smell is lacking, or even
rudimentary. All bats have a strong odor, the purpose of which is
probably to attract others of their kind. This may be taken as an
indication that smell is well developed, for otherwise the odor would
be useless.’’ The same author enumerates several instances in which
captive bats failed to locate food which could have been found only
through the sense of smell.
Taste —I can find no statements concerning a sense of taste among
bats except the very general one by Barrett-Hamilton (1911, p. 28):
“Tn captivity practically all insects are accepted except those which
are distasteful to insectivorous animals generally.’’
232 University of California Publications in Zoology [Vou. 17
Hasirs or Bars
The habits of our California bats are, unfortunately, but little
known. The time and extent of the breeding season, migration and
hibernation, the choice of diurnal retreats, and favorite feeding-
erounds, the methods of securing and devouring prey, the nature of
the food, the economic value of bats—these are only a few of the many
points on which data are as yet almost wholly lacking.
The seanty material which the writer has been able to gather
regarding the habits of our bats is given in the succeeding pages under
the subheading Natural History, at the close of the description of each
species. However, certain general topics, such as migration, are sep-
arately discussed in the paragraphs immediately following.
Migration.—Recorded observations concerning the migration of
bats are few, and in California the study of bat migration has not
progressed beyond the observation that some species which are com-
mon in fall and winter are not to be seen during the summer months,
while others known to oceur during the summer are apparently absent
during fall and winter.
Among the bats of California are found three species which occur
in suitable localities throughout the whole United States and in British
Columbia. These are the Silvery-haired Bat (Lasionycteris noctiva-
gans), the Hoary Bat (Nycteris cinercus), and the Red Bat (Nycteris
borealis), of which latter species several races occur. These three
bats are known to be migratory in at least portions of their ranges.
and some of the facts relating to their migration are of general
interest.
As Dr. C. Hart Merriam (1887, p. 85) points out, all North Amer-
ican bats, except in those places where their habits have been modified
by proximity to man, may be classed either as cave-dwelling or as
tree-dwelling, according to the places in which they spend the day.
As a rule, the cave-dwelling species live in large colonies, while the
tree-dwelling species live singly or in but small companies. Now it
is well known that the temperature in caves is little affected by the
condition of the atmosphere outside, while the temperature of holes
in trees and recesses in the foliage is about the same as that of the
surrounding air. ‘These three migratory bats are here in California
foliage-dwellers, exposed as directly as are birds to changes in atmos-
pheric temperature.
Merriam gives no details of the extent or exact northward and
1918] Grinnell: A Synopsis of the Bats of California 233
southward movements of the silvery-haired or hoary bats in the east,
but he records the occurrence of the former species about the hght-
house on Mt. Desert Rock, thirty miles off the coast of Maine, in
spring and fall. On this treeless islet bats are at other times un-
known.
Seton (1909, p. 1175-1176) states that on the Red River at Win-
nipeg he has found the silvery-haired bats common from the vernal
equinox until about September 21, after which they are neither seen
flying, nor found in their accustomed daytime haunts.
The hoary bat breeds only in the boreal zone of North America,
but it has never been recorded in this zone in winter. In winter it
occurs regularly at least as far south as the southern border of the
United States. It has been taken on the Bermudas, showing that it
is able to cross a strip of ocean having at its narrowest extent a width
of over six hundred miles. It is, however, as suggested by Miller,
much more likely that the bats commence their ocean journey at
some point much farther to the north, such as Cape Cod, the distance
from which to the Bermudas is about seven hundred miles. Of this
bat’s occurrence in the Bermudas, J. M. Jones (1884, pp. 145-146)
‘
says that it ‘“‘is observed occasionally at dusk during the autumn
months hawking about according to its nature in seareh of insects;
but as it is never seen except at that particular season it is clear that
it is not a resident, but merely blown across the ocean by those violent
northwest gales which also usually bring numbers of birds from the
American continent.’’ The latest date at which Seton has recorded
hoary bats in Toronto is mid-September. Major Mearns (in Howell,
1908, p. 37) records a diurnal flight of hoary bats at Fort Snelling,
Minnesota, but no details are given.
In regard to the red bat, Seton (1909, p. 1189) says that in sum-
mer, in Manitoba, this bat roosts in trees, it is solitary, and is not
known to frequent caves. ‘‘In winter it is known to gather in vast
numbers in the caves of its more southerly range.’’ Seton gives no
authority for the statement that this bat hibernates in caves, and I
can find no confirmation of his statement in accounts by other writers.
In southern California where the red bat is often found in evergreen
trees in winter it has never, so far as I am aware, been found in caves.
Writing from the Hudson Highlands of New York, where this bat is
very common in summer, Mearns (1898, p. 345) says: ‘‘During the
latter part of October and the first week of November, I have seen
great flights of them during the whole day. In 1876 I noted that all
234 University of California Publications in Zoology [Vov. 17
of the individuals shot from any single flock were of the same sex,
though another flock might yield all of the opposite sex.”
Rhoads (1903, p. 213) writes:
I have observed this species returning from apparently extensive flights over
the ocean on the N[ew] Jersey coast in the early morning before sunrise. On
one or two occasions in September single individuals haye been observed flying
directly toward the shore, so exhausted as to make little progress against a land
breeze and alighting on the nearest object as soon as land was reached. It is
possible that these had been blown to sea during their migrations along the coast.
It is not generally realized that bats possess powers of flight
superior to those of many birds; yet this may be inferred from the
fact that a red bat has been known to catch flies in the air while bur-
dened with young that tegether weighed more than she did.
In August and September, 1890 and 1891, Miller (1897a, pp. 541—
543) had the opportunity of watching the appearance and disappear-
ance of the above three species of bats at a locality where none could
be found during the breeding season. Highland Light, where the
observations were made, is situated near the edge of a high place in
the series of steep bluffs of glacial deposit which form the outer side
of Cape Cod, Massachusetts. The light, which is less than ten miles
from the northern extremity of the cape, is separated from the main-
land toward the east and northeast by from twenty-five to fifty miles
of water. The bluff on which it stands rises abruptly from the beach
to a height of one hundred and fifty feet. All three species (the
silvery-haired, hoary, and red bats) were found flying for the most
part along the face of the bluff, feeding upon the myriads of insects
blown there by the prevailing southwest winds. The bats were never
seen by Miller in the daytime, although he made diligent search for
them. He suggests that they may have found shelter by day in the
dense, stunted scrub-oak growth, which crowns the bluff in many
places. In 1890 the first bats of the season were seen August 21, and
the last September 12. In 1891 the first date recorded is August 25,
and the last September 13. Separate records were kept of the differ-
ent species. The numbers of bats of each species seen in a single
evening varied from one to sixteen.
An interesting observation upon the diurnal migration of bats was
made by Howell (1908, pp. 35-87) at Washington, D. C., upon the
morning of September 28, 1907. The observer first noted bats at
8 a.m. They were all flying with the wind, which was southwest.
They did not fly in flocks, but singly, usually but four or five being
1918 | Grinnell: A Synopsis of the Bats of California 235
in sight at one time. The manner of flight was quite unusual, for
instead of the erratic zigzag course commonly followed by bats when
seeking their food at dusk, the flight of those noted on this occasion
was very steady, consisting chiefly of a sailing or drifting motion,
with occasional short flappings of the wings. The height above the
ground was estimated as varying from one hundred and fifty to four
hundred feet. With the aid of a glass three different sizes were noted,
but of course it was quite impossible to identify the species. The
flight lasted for over an hour and more than one hundred individuals
were observed.
Howell suggests the possibility that such diurnal migrations are of
regular occurrence, but if that be so, it seems remarkable that they
have not been more frequently observed.
Food and Feeding Habits——There are among bats species which
eat fruit, some which eat fish, others which subsist upon blood, and
lastly, and by far the most numerous, are those species which eat only
insects. The bats native to California are, with one possible excep-
tion (see under Macrotus californicus, p. 257), strictly msectivorous.
Campbell (1913, p. 1176) states definitely that bats will eat pieces
out of hams and bacon left in smoke houses. I know of no other refer-
enee to such a habit, outside of nursery rhymes.
Most of our bats catch their prey upon the wing and devour it
without alighting. As they eat the insects they bite off and reject the
hard parts, which fall to the ground. The soft edible parts are very
finely triturated by the sharp teeth, so that it becomes quite difficult
to identify the insect remains found in the stomach of a bat. How-
ever, one California bat, Antrozous pacificus, brings at least the larger
of its insect victims to its roosting place, and hangs there while eating.
From the insect remains on the floor beneath the roost it is a simple
matter to learn the nature of the food (see p. 355).
At least two British bats (Barrett-Hamilton, 1911, p. 172) hold
the tail curved beneath them in flight, and one of them has been
observed to use the sac thus formed by the interfemoral membrane as
a pouch into which it thrusts a struggling insect until it has secured
a firm grip upon it. It would be of interest to learn whether any of
our species share this habit.
As is the case in other groups of mammals, all our bats become
very fat in autumn. Specimens secured in late winter or early spring,
on the other hand, are invariably lean. The fat stored up in the fall
is absorbed during the period of hibernation or emigration. Those of
236 University of California Publications in Zoology [Vou.17
the non-migratory species which remain on the wing throughout the
year seem to rely in good part upon such stored fat to help tide them
over the winter months, during which flying insects are available in
but seanty numbers.
Breeding Habits——I have been able to collect data which show the
number of young in most of our species of bats, and the approximate
time of birth, but the time and extent of the mating season is still
unknown. According to Barrett-Hamilton (1911, pp. 31-32), Messrs.
Rollinat and Trouessart have established the fact that in France the
normal mating time is autumn. Barrett—Hamilton says:
At that season spermatozoa are found numerously in the uterus of the adult
female, and the organs of the male are also functional. Ovulation is, however,
postponed until the termination of hibernation, during which period the ovaries
are quiescent, but the spermatozoa retain their activity in the uterus until
fertilisation takes place, in April. On this point all authorities are agreed, and
the facts, subject to correction as to details, may be taken as substantiated. An
alternative view, that ovulation and fertilisation may take place in the autumn and
winter, with subsequent postponement of the development of the embryo, seems
to be unsupported by facts.
Recorded observations of autumnal mating in certain species of the
eastern portion of the United States incline one to the belief that
nearly related races occurring in California may possess similar habits.
Murphy and Nichols (1913, p. 11), writing upon the bats of Long
Island, confirm earlier reports of autumnal mating in the eastern red
bat (Nycteris borealis borealis).
VOICE
As is well known to all naturalists, the voices of bats are high and
shrill; in other words, the vibrations are of high frequency. Barrett-
Hamilton (1911, p. 43) states that a young bat calls for its mother as
persistently as any other young animal, and in commenting upon the
shrillness of the voices of bats suggests that some of their cries are
pitched on a higher seale than that to which any human ear is attuned.
At least two gregarious bats, Antrozous and Nyctinomus, are quite
noisy in their daytime haunts, squeaking almost continually as they
jostle one another restlessly. Coues (1867, p. 284) mentions the
squeaking and scratching of Antrozous pallidus in the chinks of the
officers’ quarters at Fort Yuma. When sleeping out-of-doors near
a loft oceupied by a colony of Pacifie pallid bats, the present writer
has often heard their shrill voices far into the summer night.
1918 | Grinnell: A Synopsis of the Bats of California 237
ENeEmigs oF Bats
It is axiomatic that the birth-rate of a species is indicative of the
relative degree of danger to which its individuals are subject. Among
bats the usual number of young is but one or two at a birth, and but
one litter is produced annually. Hence it may be taken for granted
that bats have but comparatively few enemies. Three reasons may
be advaneed to account for this comparative immunity from enemies,
namely: the probably unpalatable flavor of bats, as suggested by their
odor; the comparative safety and remoteness of their diurnal retreats ;
and the fact that they are both volant and crepuseular. In this con-
nection it may be observed that a comparatively slow rate of repro-
duction is to be found also among nighthawks and owls, which like-
wise fly at twilight or nocturnally.
C. H. Merriam (in Murphy and Nichols, 1913, p. 10) says: ‘‘ Bats
have been found in both owl and hawk pellets a number of times, but
only rarely in hawks’. They have been found also in the stomachs
of large trout, and it goes without saying that they are sometimes
discovered and eaten by some of the smaller predatory carnivores.’
Seton (1909, p. 1181) says: ‘‘At Chilliwack Lake, in British
Columbia, the rainbow trout are of great size, eight pounds to twelve
pounds, and these giants were often seen by Professor John Macoun,
leaping after the bats that skim the surface of the lake at evening.
In one ease he thinks he saw a bat captured by the trout, and is satis-
fied that the fish would not jump so persistently if they did not fre-
quently sueceed.’’ This is of interest, but obviously inconclusive.
Fisher (1893) records the results of the examination of the
stomach contents of 2690 hawks and owls inhabiting the United States,
including forty-nine species and subspecies. In this list (p. 180)
we find but one mention of the bat, a single silvery-haired bat
(Lasionycteris) that was found in the stomach of a great horned
owl. Altum (1863, pp. 48, 217) records the examination of 703 pellets
disgorged by European barn owls, among which were found a total of
2551 skulls, sixteen being of bats.
Barrows (1884, p. 29), in a paper on birds of the Lower Uruguay
River, South America, states that at night the deserted corridors of
the college at Concepcion was one of the barn owl’s favorite hunting-
grounds for bats. Bailey (1905, p. 211) states that he found two
lower jaws of the large brown bat in pellets under the nest of a great
horned owl in Texas. Miller (1904, p. 337), in an article on Cuban
bats, mentions an individual barn owl which fed largely on bats.
238 University of California Publications in Zoology [Vou.17
It has been stated to the author that house cats find the bodies of
bats unpalatable. No opportunity has been found for experimen-
tation along this line.
It is clear that the enemies of bats are few; only one seems to
make more than slight inroads upon their numbers. This enemy is
man, who is often guilty of wholesale destruction of individuals of
eregarious species. Where colonies of bats have taken possession of
attics, barns, or church steeples they are often ruthlessly destroyed
when once their squeaking, or the disagreeable odor emanating from
their haunt, has betrayed them. A simple method of disposing of a
colony of bats which has proved a nuisance is to board up, or sereen
over, all the entrances to such retreats in the evening when the occu-
pants are on the wing.
Economic VALUE OF BATS AND THEIR CONSERVATION
Many detailed studies have been made and much written to show
the value of our insectivorous birds. As much more might be written
to show the economic value of our bats. For the latter are undoubt-
edly as important in keeping in check crepuscular and nocturnal
insects as the birds are in destroying day-flying species. Among the
insects destroyed there are not only many species which are harmful
to agriculture, but also disease-carrying inseets, such as the mosquito
hosts of malaria.
Dr. C. A. R. Campbell (1918, pp. 1175-1181) has estimated that
at Mitchell’s Lake, Texas, 90 per cent of the food of bats consists of
malaria-carrying mosquitoes. At this place Dr. Campbell built a
bat roost in 1911, and during the succeeding two years, when the roost
had become populously tenanted, he not only found that the number
of mosquitoes had decreased materially but also that malaria had
become much less prevalent in the region. In the neighborhood where
these experiments were carried on not only were the people much
annoyed by mosquitoes before the erection of the roost, but stock also
suffered severely, horses and cattle becoming thin and anemic in spite
of being well fed. Campbell (1918, p. 1176) believes that the bat
itself is protected against the depredations of the mosquito by the
peculiar formation of the hairs, which in the bat are not smooth and
round, but in appearance likened to a number of morning-glory
flowers strung on a straw (fig. A).
Campbell suggests the wisdom of erecting roosts, similar to the one
1918 | Grinnell: A Synopsis of the Bats of California 239
which he has designed, in all parts of the country where mosquitoes
are prevalent. He points out that in time the cost of these roosts
is defrayed through the value of the guano which may be collected
for fertilizer and which has, according to statements submitted by
him, a commercial value of about thirty dollars per ton.
It is interesting to note that in at least one city of the United
States, San Antonio, Texas, bats are now protected by law. This city,
in June, 1914, passed an ordinance to prohibit the destruction of bats
within the city limits, and preseribed that anyone violating the pro-
visions of the ordinance should be fined not less than five dollars, or
more than two hundred dollars, for each bat so killed.
nay Nee f
a b c d
Fig. A. Portion of a hair from the dorsum of Nyctinomus mexicanus, no. 19092.
X 288.
Fig. B. Portion of a hair from the dorsum of Humops californicus, no, 4326.
X 288.
Fig. C. Portion of a hair from the dorsum of Nycteris cinerea, no. 20778.
X 288.
Fig. D. Portion of a hair from the dorsum of Myotis longicrus interior, no, 17790.
X 288.
The earlest published record which | have found of the protection
of bats because of their usefulness to man is that of J. K. Townsend
(1839, p. 325), who says that the great-eared bat (Corynorhinus)
Inhabits the Columbia river district [Washington], rather common. Frequents
the store houses attached to the forts, seldom emerging from them even at night.
This, and a species of Verpertilio [sic], (V. subulatus,) which is even more
numerous, are protected by the gentlemen of the Hudson’s Bay Company, for
their services in destroying the dermestes which abound in their fur establish-
ments.
240 University of California Publications in Zoology [Vov.17
ORIGIN OF CALIFORNIA Bats
The bones of bats are so small and delicate that they are very easily
destroyed, and it is probably for this reason that they are compara-
tively seldom found as fossils. In only two instances known to the
writer have fossil bats been discovered in California. The palaeonto-
logical collection of the University of California contains a skull which
I have identified as Myotis longicrus, from Samwell Cave, Shasta
County ; and in the same collection is a broken skull and several lower
jaws of Antrozous pallidus from Potter Creek Cave, also in Shasta
County. In both cases the horizon is doubtfully Pleistocene.
Gregory (1910, p. 319) says of the origin of the bats:
The group was very highly specialized as far back as the Eocene and the
palaeontological evidence as to its origin is therefore of a negative character, as
in the case of the Rodents, Edentates and all other groups which probably acquired
many of their ordinal characters before the known Tertiary record began.
Nevertheless there can be little doubt that the Chiroptera are an offshoot of late
Mesozoic or early Tertiary arboreal Insectivores, which must have resembled
Galeopithecus in many characters.
It is of interest to note the present world-distribution of the eleven
genera of bats found within the State of California. The ranges
here given are from Miller (1907). Four genera occur in both the
Old World and the New. These are: Myotis, found in both hemi-
spheres to the northern and southern limits of tree growth; Pipistrel-
lus, found in the Eastern Hemisphere to the limits of tree growth, but
in the Western Hemisphere confined to North America from the
northern United States (except in the Boreal zone) to southern Mex-
ico; Nyctinomus, found in the warmer portions of both the Eastern
and the Western hemispheres; and EH ptesicus, found in Afriea, Mada-
gascar, Australia, Asia (except the Malay region), and America from
southern Canada southward (exeept Lesser Antilles).
Of the seven genera confined to the New World, Nycteris is the
most widely distributed, occurring in North America northward to
the limit of tree growth and southward to include the Bahama Islands
and Greater Antilles, as well as the Galapagos and Hawaiian Islands.
Lasionycteris occupies northern North America, south through the
United States. Antrozous is found in western North America from the
Columbia River to central Mexico. Corynorhinus occurs throughout
the warmer portions of North America to southern Mexico. Macrotus
occurs in the warmer parts of North America from Guatemala to
1918 | Grinnell: A Synopsis of the Bats of Califorma 241
southern California and Arizona, also in the Greater Antilles and the
Bahama Islands. EHuwmops is found in the warmer parts of America
north to the southwestern United States, and in the Greater Antilles.
Euderma is the most closely restricted of our genera, having so far
been found only in the southwestern United States.
The genera of bats now occupying California may be inferred to
have reached their present location from three different sources.
First, cosmopolitan genera, namely Myotis, Eptesicus, Nyctinomus
and Pipistrellus probably originated in the Old World. Second,
Nycteris, Lasionycteris, Antrozous and Corynorhinus probably orig-
inated in temperate North America. Third, Macrotus and Humops,
southern genera, probably originated in tropical America and
now reach their northernmost limit along the southern edge of our
state. With the second group may be classed the genus Huderma,
now confined to the extreme southwestern United States, and most
closely related to the genera Corynorhinus and Plecotus, of the tem-
perate zones.
GEOGRAPHIC DISTRIBUTION OF Bats
Bats occur throughout the eastern and western hemispheres to
the northern and southern limits of tree growth (Miller, 1907, p. 43).
Their day-time haunts vary from the cool, humid depths of mountain
caves or tunnels to the torrid and arid rock-crannies of the desert.
Their nocturnal foragings must involve extremes of temperature and
humidity equally as great. When each species is considered sep-
arately, however, we find it to be nearly, if not quite, as strictly
limited in range as are other mammals, and this in spite of better
powers of locomotion.
It has been so generally supposed that bats show far less geo-
eraphie variation than do other small mammals that the opinion of
two recent workers in the order Chiroptera may properly be quoted
here. One of these, Barrett-Hamilton (1910, p. 20), says:
Neglect of bats is a grave error in studying geographical distribution, since,
inasmuch as these creatures are possessed of the power of surmounting obstacles
which to other mammals must be insuperable, their permanent restriction to definite
regions must be due to causes of fundamental importance. And, whereas the
wings of bats should have enabled them to occupy with uniformity the entire
extent of the British Islands, we find in fact that their distribution therein is not
less restricted than that of other mammals.
242 University of California Publications in Zoology [Vov.17
The second authority, Andersen (1912, p. Ixxvi), observes:
The evidence afforded by the geographical distribution of Bats has generally
been considered of doubtful value; hence they have either been entirely excluded
from the material worked out by zoogeographers or at least treated with pro-
nounced suspicion, as likely to be more or less unreliable documents of evidence.
This unwillingness or hesitation to place Bats on an equal zoogeographical footing
with non-flying Mammalia would seem to be due, partly to the preconceived idea
that owing to their power of flight Bats must evidently have been able easily to
spread across barriers which, in ordinary circumstances, are insuperable for
wingless Mammalia; partly to the fact that hitherto very often whole series of
distinct forms have been concealed under one technical name. So long as (to
mention only three cases among many) ‘‘ Macroglossus minimus’’ was believed to
range unchanged from the Himalayas to New Guinea, Australia, and the Solomon
Islands (now two distinct genera, thirteen recognizable forms), or ‘‘Cynopterus
marginatus’’ over India, Ceylon, Indo-China, and Indo-Malaya (now six species,
fourteen forms), or ‘‘ Rhinolophus ferrum-equinum’’ uniformly over Europe, Asia,
and Africa (now numerous distinet forms), they were undoubtedly of questionable
value as zoogeographical material. But these and similar anomalies invariably
disappear as soon as modern methods of discrimination applied on vastly in-
creased material render it possible to draw the lines of separation between the
species (and their local modifications) somewhat more closely in accordance with
the lines drawn by Nature. The second argument referred to above, that the
spreading of Bats from one locality to another must obviously have been greatly
facilitated by their possession of wings, may in theory appear plausible enough,
but when tested on the actual distribution of the species and subspecies it proves
to be of much less importance than commonly supposed; it rests, in reality, on
a confusion of two different things: the power of flight no doubt would enable
a Bat to spread over a much larger area than non-flying Mammalia, but, as a
matter of fact, only in very few cases is there any reason to believe that it has
caused it to do so.
Matthew (1915, p. 227) remarks with regard to this paragraph:
The belief that bats are more easily able to cross ocean barriers than non-
flying mammals is probably based, not on the preconceived idea that they could,
but upon the plain fact that they have done so far more frequently. Birds and
bats are found upon numerous oceanic islands where no non-flying mammals,
and very few non-flying animals at all, exist. That they have wings and occa-
sionally use them for so long a journey, whether voluntarily or involuntarily, is
a natural explanation. I cannot see any other reasonable interpretation of the
fact that they are present and the terrestrial mammals absent in so many remote
oceanic islands. With bats, as with most birds, the intervening ocean acts as a
hindrance, but their wider distribution shows that it is less of a hindrance than
with terrestrial mammals.
Matthew’s comments are good, so far as they go, but it seems to
the present writer that he has missed the main point of the thesis,
which is, that conditions of temperature and humidity limit the dis-
tribution of bats as strictly as they do that of other groups of mam-
mals. Of course, given a chain of islands of identical climatic con-
ditions, the distribution of bats would without doubt be more uniform
1918] Grinnell: A Synopsis of the Bats of California 243
than that of non-volant mammals; but were each of these islands to
exhibit a unique condition of temperature and humidity, it may be
ventured that the bats inhabiting them would be found to represent
distinct races, Just as do birds in similar eases.
In California there is not a single obvious barrier to the distri-
bution of any species of bat; yet not one of the thirty-one forms
inhabiting the state has been found to be distributed uniformly
throughout the entire area. In the most abundantly represented
genus, Myotis, at least fourteen forms are to be recognized, each
strictly limited in the breeding season to a certain zonal and faunal
area. To illustrate the point: Myotis orinomus has never been taken
outside of the semi-arid region of the higher portion of the Upper
Sonoran zone.
DENTITION
Young bats are born with a complete milk-dentition, the function
of which is probably to enable them to hold on to their mothers. These
teeth differ both in number and form from those cf the permanent set.
They are homodont, slender, and sharply recurved, and resemble the
teeth of seals and cetaceans more nearly than those of adult bats
(Barrett-Hamilton, 1910, p. 15).
The permanent teeth consist, as in other mammals, of four kinds:
incisors, canines, premolars, and molars. These teeth differ widely
among the different genera and species, in both numbers and form,
there being, for example, but twenty teeth in the jaws of the blood-
sucking genus Desmodus, and these so specialized that it would be
impossible for the animal to masticate; while the highest number of
teeth present in any known bat is thirty-eight (Miller, 1907, p. 23),
the number present in the genus Myotis.
In several of the species here listed some of the teeth are so minute
as to be functionless and are difficult to find; in fact they sometimes
drop out of the jaw. In Nyctinomus mexicanus, for instance, the
number of incisors appears to be variable; and in Myotis occultus
the middle upper premolar, normally present among related species,
is lacking in half the known specimens.
CoLORATION
In common with other nocturnal or crepuscular mammals bats
have, as a rule, a much more sober coloring than diurnal mammals,
244 University of California Publications in Zoology [Vou.17
birds or reptiles, being for the most part of somber grays or browns,
with the ventral surface a lighter tint of the same color that pervades
the back. Among exotic bats there are notable exceptions to this
rule, but among our Californian species only two diverge in color from
the quiet tones of their fellows. These are the spotted bat (Huderma
maculatum), which is black on the dorsal surface, with three large
white patches; and the western red bat (Nycteris borealis teliotis),
which varies in color from bright rufous-red or fawn to yellowish gray.
Young bats, while of the same general color as adults of the same
species, are usually darker and duller than their parents.
All color names used in the following pages are taken from Ridg-
way’s Color Standards and Color Nomenclature (1912).
AGE VARIATION
In determining the age of bats the writer has followed the sugges-
tion of Miller (1897b, pp. 8-9), who finds that in adults the finger
joints are small and compact, the epiphyses no longer visible, and the
phalanges of essentially the same diameter throughout; whereas in
young specimens the joints are ‘‘large and loosely formed, with
epiphyses separate from the ends of the phalanges and metacarpals,
both of which are distinctly enlarged for some distance from the
joint.’’
The matter of determining the age of individuals is important
because of the fact that young bats, even when nearly full grown,
present characters different enough from those of the adults to cause
confusion in identification. In general, the fur of immature speci-
mens is shorter and more woolly than that of adults, and the color is
darker and duller. The young of Lasionycteris, however, are a notable
exception to this rule, the pelage of young examples being long and
silky, and the hairs having beautiful silvery tippings, which in the
adults are less perfectly shown.
SEXUAL VARIATION
There is but little sexual variation among North American bats,
and in those inhabiting California I have found such as is present to
consist only of a slightly greater average size of the females in a
few of the species, for example, in Pipistrellus.
1918] Grinnell: A Synopsis of the Bats of California 245
NOMENCLATURE
The nomenclature here used is for the most part that adopted by
Miller (1912).. Where it has seemed necessary to deviate from this
standard, an explanation is given in the account of the species con-
cerned under the heading Synonymy and History.
CLASSIFICATION
All bats are included in the order Chiroptera and are in common
distinguished by the following characters:
Mammals with the front limbs modified for true flight, the fingers greatly
elongated (the third usually at least equal to head and body) and joined together
by a membrane which extends to sides of body and legs; shoulder girdle much
more developed than pelvis, the sternum usually keeled; knee directed backward
owing to rotation of leg for support of wing membrane (Miller, 1907, p. 43).
The families of bats fall naturally into two main groups, the
Megachiroptera and the Microchiroptera. The first of these groups,
the Megachiroptera, or fruit-eating bats, represents an evolutionary
stage much nearer to ordinary mammals than the latter. Its mem-
bers are distinguished externally by the presence of a claw on the
end of the second digit as well as on the pollex, by the completeness
of the ring formed by the margin of the ear, and by the absence of a
tragus. Andersen (1912, p. vi), the most recent worker on the group,
recognizes 228 genera. These are confined to the tropical and sub-
tropical regions of the Old World, east to Australia, Samoa, and the
Caroline Islands. Hence none of them fall within the scope of the
present paper. The second group, the Microchiroptera, comprises
those bats characterized by the absence of a claw on the second digit,
by the incompleteness of the ring formed by the margin of the ear,
and by the presence, normally, of a tragus (Miller, 1907, p. 44). The
same author (loc. cit., p. 78) recognizes six hundred genera which he
has grouped into sixteen families. The Microchiroptera are found in
both the eastern and western hemispheres.
Only three of the families listed by Miller occur in the Western
Hemisphere north of the Bahama Islands and Central Mexico. These
are: the Phyllostomidae, represented in California by a single species;
the Vespertilionidae, represented by twenty-six species and subspecies ;
and the Molossidae, represented by four species. These bats may
be identified with the aid of the appended keys, which are wholly
artificial and make no pretense of indicating natural relationships.
Anyone interested in the classification of the families and genera
of bats by means of skeletal and dental characters is referred to
Miller’s (1907) The Families and Genera of Bats.
246 University of California Publications in Zoology [Vou. 17
Thumb
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Free edge of interfemoral membrane
Fig. E. Wing of Hptesicus fuscus (drawn semidiagrammatically from spect-
men no. 23522), X 1.11; labeled to show names of parts, as used in the present
paper.
KEY TO CALIFORNIA BATS
I. Basep oN EXTERNAL CHARACTERS
PAGE
1. With noseleaf (see text-fig. F). Macrotus californicus 252
1’. Without noseleaf.
2. Tail extending conspicuously beyond free edge of interfemoral mem-
brane (see pl. 19). /
3. Total length greater than 140 millimeters; no horny exerescences
on anterior margin of ear.
Eumops californicus 370
3’. Total length less than 140 millimeters; a row of horny excrescences
on anterior margin of ear (see text-fig. W).
4. A fold of membrane extending from the inner third of the femur
to the middle of the tibia, forming a pocket at the thigh.
Nyctinomus femorosaccus 360
1918 | Grinnell: A Synopsis of the Bats of California
247
PAGE
4’. No fold of membrane extending from the inner third of the
femur to the middle of the tibia.
5. Length of forearm more than 50 millinreters.
Nyctinomus depressus :‘
5’. Length of forearm less than 50 millimeters.
Nyctinomus mexicanus 3
2’. Tail extending at least to edge of interfemoral membrane, but never
much beyond (see pl. 18).
6. Nostrils opening forward beneath a conspicuous horseshoe-shaped
ridge (see text-fig. U).
7. Length of forearm more than 52 millimeters; color of back yel-
lowish drab brown; habitat west of the desert divides.
Antrozous pacificus 3
7’. Length of forearm less than 52 millimeters; color of back whitish
drab-gray; habitat east of the desert divides.
Antrozous pallidus
6’. Nostrils not opening beneath a horseshoe-shaped ridge.
8. Dorsal surface of interfemoral membrane furred, at least on
basal half (see pl. 17, fig. 10).
9. Total length greater than 120 millimeters. | Nycteris cinerea
9’. Total length less than 120 millimeters.
10. General color ranging from rufous red or fawn to yellowish
gray. Nycteris borealis teliotis
10’. General color blackish chocolate. Lasionycteris noctivagans
8’. Dorsal surface of interfemoral membrane naked except for scat-
tering hairs, or furred only at extreme base.
11. Height of ear from crown more than 25 millimeters.
12. Tragus broadly rounded at tip; back blackish with three
white patches (see pl. 16, fig. 9). Euderma maculatum 3:
12’. Tragus slender, pointed at tip (see text-fig. S); color not
blackish and no white markings present.
13. General color wood brown; habitat the arid portions of
the Upper and Lower Sonoran zones in southern
California. Corynorhinus rafinesquii pallescens
13’. General color, natal brown; habitat the Upper Sonoran
zone in western California.
Corynorhinus rafinesquii intermedius :
11’. Height of ear from crown less than 25 millimeters.
14. Tragus short, blunt, and curved forwards (see text-fig. L).
15. General aspect of fur pale grayish; habitat Sonoran zones
east of the desert divides.
' Pipistrellus hesperus hesperus
15’. General aspect of fur pale brownish; habitat Sonoran
zones west of the desert divides.
Pipistrellus hesperus merriami
14’. Tragus not curved forwards, slender.
16. Total length more than 105 millimeters. Eptesicus fuscus
16’. Total length less than 105 millimeters.
17. Ear when laid forward extending 7 to 10 millimeters
beyond nostril. Myotis evotis
348
330
323
300
os
a
340
305
311
314
291
248
University of California Publications in Zoology (Vou. 17
PAGE
17’. Ear when laid forward extending less than 6 milli-
meters beyond nostril.
18. Free border of uropatagium distinctly fringed with
fine hairs. Myotis thysanodes
18’. Free border of uropatagium faintly or not at all
fringed.
19. Length of tibia 15 to 19 millimeters.
20. Length of tibia less than 17 millimeters.
Color of back, drab; habitat Lower Sonoran
zone east of the desert divides.
Myotis velifer *
Color of back, light brown; habitat Boreal
zone on mountains of east-central and north-
ern California. Myotis lucifugus altipetens
Color of back, dark brown; habitat extreme
northern coast belt.
Myotis lucifugus alascensis
20’. Length of tibia more than 17 millimeters.
Color of back, yellowish brown; habitat south-
ern Sierra Nevada and mountains of south-
ern California. Myotis longicrus interior *
Color of back, dark brown; habitat central and
northern Sierra Nevada and humid coast
belt. Myotis longicrus longicrus *
19’. Length of tibia less than 15 millimeters.
21. Foot 5 to 6 millimeters in length.
Color of back, pale grayish brown; habitat
Lower Sonoran zone east of the desert di-
vides. Myotis californicus pallidus
Color of back, bright brown; habitat Upper
Sonoran and Transition zones south of the
36th parallel. Myotis californicus quercinus :‘
Color of back, dark brown; habitat Upper
Sonoran and Transition zones north of the
36th parallel. Myotis californicus californicus *
21’. Foot 7 to 10 millimeters in length.
22. Height of tragus more than 6.5 millimeters.
Myotis orinomus
22’, Height of tragus less than 6.5 millimeters.
Color of back, cinnamon brown; habitat val-
ley of the Colorado River.
Myotis occultus
Color of back, light buff; habitat Lower
Sonoran zone east of the desert divides.
Myotis yumanensis yumanensis
Color of back, buffy brown; habitat Sonoran
and Transition zones west of the desert
divides. Myotis yumanensis sociabilis :
Color of back, dark brown; habitat extreme
northern coast belt.
Myotis yumanensis saturatus *
2907
263
267
288
290
261
278
1918] Grinnell: A Synopsis of the Bats of California 249
II. BASED ON SKULLS AND TEETH
1. Total number of upper incisors 2. PAGE
2. Greatest length of skull 30 millimeters or more; upper incisors diverg-
ing at tips (see pl. 20, fig. 22). * Eumops californicus 370
2’. Greatest length of skull less than 25 millimeters; upper incisors con-
verging at tips.
3. Four teeth in upper jaw behind canine, none minute.
4. Greatest length of skull 19-21 millimeters. Antrozous pallidus 348
4’. Greatest length of skull 22-24 millimeters. Antrozous pacificus 352
3’. Five teeth in upper jaw behind canine, first one minute.
5. Rostrum louger than wide.
6. Greatest length of skull more than 22 millimeters.
Nyctinomus depressus 357
6’. Greatest length of skull less than 22 millimeters.
7. Greatest length of skull more than 18 millimeters.
Nyctinomus femorosaccus 36)
7’. Greatest length of skull less than 18 millimeters.
j Nyctinomus mexicanus 361
od’. Rostrum wider than long.
8. Greatest length of skull more than 15 millimeters.
Nycteris cinerea 330
8’. Greatest length of skull less than 15 millimeters.
Nycteris borealis teliotis 325
1’. Total number of upper incisors 4.
9. Upper incisors completely filling space between canines.
Macrotus californicus 252
9’. Upper incisors not completely filling space between canines.
10. Total number of teeth 32. Eptesicus fuscus 314
10’. Total number of teeth 34 to 38.
11. Total number of teeth 34.
12. Greatest length of skull less than 15 millimeters.
Pipistrellus hesperus hesperus 305
Pipistrellus hesperus merriami 311
12’. Greatest length of skull more than 15 millimeters.
Euderma maculatum 336
11’. Total number of teeth 36 or 38.
13. Total number of teeth 36.
14. Sagittal and occipital crests well defined (pl. 21, fig. 30,
pl. 22, fig. 42). Myotis occultus 261
14’. Sagittal and occipital crests not well defined.
15. Upper surface of rostrum with distinct depression on each
side between lachrymal region and external nares.
Lasionycteris noctivagans 300
15’. Upper surface of rostrum with no depression on each
side between lachrymal region and external nares.
Corynorhinus rafinesquii pallescens 540
Corynorhinus rafinesquii intermedius 344
University of California Publications in Zoology [Vo
13’. Total number of teeth 38.
16. Greatest length of skull 16 millimeters or more.
17. Sagittal crest well defined; rostrum broad (pl. 21, fig. 32).
Myotis velifer
17’. Sagittal crest not well defined; rostrum slender.
18. Frontal region conspicuously inflated (pl. 21, fig. 35;
and pl. 22, fig. 47); zygomatic breadth more than
9.5 millimeters. Myotis thysanodes
18’. Frontal region not conspicuously inflated (pl. 21, fig.
34, and pl. 22, fig. 46); zygomatic breadth less than
9.5 millimeters. Myotis evotis
16’. Greatest length of skull less than 16 millimeters.
19. Sagittal and occipital crests well defined (pl. 21, fig. 30,
and pl. 22, fig. 42). Myotis occultus
19’. Sagittal and occipital crests not well defined.
20. Dorsal profile of skull distinctly flattened (pl. 22, fig.
38). Myotis orinomus
20’. Dorsal profile of skull not flattened.
21. Occipital elevation distinct (pl. 22, fig. 40).
Myotis longicrus longicrus
Myotis longicrus interior
21’. Occipital elevation not distinct.
22. Interorbital constriction less than 3.5 millimeters.
Myotis californicus californicus
Myotis californicus quercinus
Myotis californicus pallidus 2
22’, Interorbital constriction more than 3.5 millimeters.
23. Zygomatie breadth more than 8.5 millimeters;
greatest length of skull more than 14 milli-
meters.
Myotis lucifugus altipetens
Myotis lucifugus alascensis 2
N
ygomatic breadth less than 8.5 millimeters;
greatest length of skull usually less than 14
millimeters.
Myotis yumanensis yumanensis
Myotis yumanensis sociabilis
Myotis yumanensis saturatus
L. 17
PAGE
259
297
291
261
290
1918 | Grinnell: A Synopsis of the Bats of California
251
AVERAGE MEASUREMENTS OF THE CALIFORNIA SPECIES OF BATS
ARRANGED IN SEQUENCE FROM SMALL TO LARGE ACCORDING To ToTAL LENGTHS
Species
Pipistrellus hesperus hes-
TPEVSWS coset tee Staccens eaceereee
Pipistrellus hesperus mer-
riami a
Myotis californicus eali-
TOT AWICWED conctabenenteee meer eeeeS
Myotis californicus pal-
UGLY cee reenereeeeeeeereres eee ee
(HUTT) eet eee
iligmpeene tee! ER eo
Myotis yumanensis yu-
HUD TTGTAGNS) Gelecen esa eeepee
Myotis yumanensis sat-
PIU UES Meee cer Seneee -ssneneocsas
Myotis orinomus ..............
Myotis lucifugus — alas-
CBUISIS) oes eno-Stececeecteceececoasas
Myotis lucifugus alti-
TOMEI) Se pa eee eee
Myotis thysanodes ...........
Myotis oecultus)-..........-...
Iv ObISH eC VOUIS) 2/22 ons
All Measurements in Millimeters
Number of
specimens
10 dig, 10 99
6 $d, 10 99
499
299
599
19
Nyetinomus mexicanus .... 10 dg, 10 92
Myotis longicrus longi-
(OTHE) eee Ree eae
Myotis velifer ...........2....:.
Corynorhinus rafinesquii
AUHESCONS |) cess2 cman sens
Macrotus californicus ......
Myotis longicrus interior
Lasionycteris noctivagans
Corynorhinus rafinesquii
intermedius ....................
Nyctinomus femorosaccus
Nyeteris borealis teliotis
Antrozous pallidus ..........
Euderma maculatum ......
Eptesicus fuseus ..............
Antrozous pacificus -.......
Nyctinomus depressus -...
Nycteris cinerea ..............
Eumops californicus ........
8 gd,
433;
3 3d;
12 3d,
1 g2
73d;
3 dd;
299
18 9°
899
929
899
499
5 29
5 Qo
ue
5 99
799
Total
length vertebrae
68.6
97.2
97.3
98.9
102.0
102.0
103.0
104.5
109.3
111.5
112.4
119.0
131.0
135.1
167.1
Tail
Tibia
Foot
10.4
8.5
11.6
7.6
9.5
12.9
13.0
Hale
16.8
Greatest
length of
Forearm cranium
28.8
28.0
35.8
41.2
35.6
37.5
41.3
37.2
40.4
39.9
49.9
38.0
39.2
42.0
47.0
38.7
49.2
50.8
45.1
55.3
60.0
51.8
72.0
‘Miller (1897b, p. 83); * Merriam (1889a, p. 23); *J. A. Allen (1891b, p. 196); 4 Ward
(1891, pp. 747-750).
252 University of California Publications in Zoology [Vou.17
Order CHIROPTERA
Suborder MicrocHIROPTERA
Family PHYLLOSTOMIDAE
This family, as represented in North America, includes bats having
a simple noseleaf, a tragus, toes (except hallux) with three phalanges
each, thumb and foot without sucking disk, and upper incisors not
separated by a space in the middle. The only genus which has been
found to occur in California is Macrotus, which is here represented
by a single species.
Genus Macrotus Gray
mee leh 2-2 gee 34
59’ © yay? PM g7g> Mang = 34.
Skull with rostrum tapering, and distinctly flattened above; auditory
bullae conspicuously enlarged, covering almost the entire cochleae,
their greatest diameter being distinctly greater than the width of the
space between them. The extremity of the tail projects beyond the
hinder edge of the broad interfemoral membrane.
Diagnosis.—Dental formula: i
Macrotus californicus Baird
California Leaf-nosed Bat
Macrotus Californicus Baird (1858, pp. 116-117). Original description;
type locality Fort Yuma, Imperial County, California.
Macerotus californicus, Baird (1859, II, p. 4, pl. 61, fig. 2). Description;
listed from Fort Yuma.
Maerctus californicus, H. Allen (1864, pp. 3-5, figs. 2-4). Description.
Macrotus californicus, Cooper (1868, p. 5). Distribution.
Macrotus californicus, Cooper (in Cronise, 1868, p. 442). Description;
occurrence near Fort Yuma. :
Macrotus waterhousii, Coues and Yarrow (1875, pp. 80-81).
Macrotus waterhousii, Dobson (1878, pp. 464-467), part. Description;
occurrence in California; habits.
Macrotus Waterhousei, True (1885, p. 604), part. Range.
Macrotus waterhousei, Bryant (1891a, p. 359). Nominal.
Macrotus californicus, H. Allen (1894, pp. 34-48, pls. 1-2). Deseription ;
listed from Fort Yuma.
Macrotus californicus, Elliot (1901, pp. 420-421, fig. 93). Diagnosis.
Otopterus californicus, Miller and Rehn (1901, p. 278). Type locality.
Macrotus californicus, Rehn (1904, pp. 441-444). Revision of genus; dis-
tribution of M. californicus.
Otopterus californicus, Elliot (1904b, pp. 653-654). Description; distri-
bution.
Otopterus californicus, Elliot (1905, pp. 509-510). General distribution;
type locality.
1918 | Grinnell: A Synopsis of the Bats of California 253
Otopterus californicus, Stephens (1906, pp. 276-277, 1 fig.). Description;
distribution ; habits.
Otopterus californicus, Lyon and Osgood (1909, p. 290). Location of type.
Macrotus californicus, Miller (1912, p. 36). Type locality; range.
Macrotus californicus, J. Grinnell (1913b, p. 275). Range in California.
Diagnosis —Size medium (total length 93-103 millimeters) ; ear
reaching beyond tip of muzzle when laid forward; distinct leaf-like
vertical appendage on nose; color, above, pale drab to broccoli brown,
below, drab, the longer hairs tipped with white.
Description: Head.—-Elongate, slender, rostrum rather attenuate ;
ears oval and slightly hairy, connected at their bases by an incised
transverse membrane, when stretched forward exceeding muzzle by
one-third entire length of ear; tragus lanceolate, one-third height of
ear. Hye almond-shaped. Nose-leaf blunt-lanceolate, higher than
width of pad from which it rises (text-figs. F, G@). Nostrils crescentie,
placed obliquely in nose pad. A triangular pad at apex of chin.
Fig. F. Side view of head of Macrotus californicus (drawn from specimen
no. 19127), X 1.00, showing simple nose-leaf, long ears, and erect pointed tragus.
Fig. G. Front view of head of Macrotus californicus (drawn from specimen
no. 19127), X 1.00, showing position of nostrils in pad at base of nose-leaf,
incised transverse membrane connecting bases of ears, and triangular pad at
apex of chin.
Limbs and Membranes.—Forearm slightly bowed. Thumb, ex-
clusive of claw, about 7 millimeters in length, and slender. Wing
short and stout; length of fifth metacarpal equaling and sometimes
exceeding that of third metacarpal. Toes compressed, giving foot a
slender aspect. The tail projects beyond edge of interfemoral mem-
brane for the length of one or one and one-half vertebrae. The mem-
branes of this bat are parehment-like rather than leathery, and are
not furred.
Pelage——F ur on body silky; on back about 8 millimeters in length,
on under surface about 6 millimeters; ear with posterior base covered
with hair of a woolly texture. Interior surface of ear, and proximal
three-fourths of anterior border of ear with scattered long hairs.
Color.—In dried specimens the color of the distal third of the hairs
on the upper surface varies from pale drab to broccoli brown. The
proximal two-thirds is white, and this shows throngh the darker outer
254 University of California Publications in Zoology (Vou. 17
color in irregular patches, due to the parting of the hairs. Fur upon
bases of ears grizzled gray throughout. Upon the lower surface of
the body the bases of the hairs are white, the distal third drab, tipped
with white, this giving a frosted appearance. The membranes in the
dried skin are clove brown. In specimens preserved in alcohol the
colors are essentially the same, save that the ventral surface lacks the
frosted appearance characteristic of dried skins, and the white bases
of the hairs are yellowed by the absorption of fat from the body.
Skull.—Rostrum at orbits twice as wide as at canines (pl. 23, fig.
54). Auditory bulla conspicuously enlarged, covering almost entire
cochlea (pl. 24, fig. 62), its greatest diameter distinctly greater than
width of space between the two bullae.
22, @il=il 2=2 38) :
59 jay’ PM ag M39 =34. Upper
incisors conspicuously unequal in size, and completely bridging space
between canines; inner pair large, in contact distally, but with roots
widely separated; lateral pair small, not extending beyond cingula of
inner pair. Lower incisors small and simple, forming a continuous
row between canines. Canines strong and simple, with well-developed
cingula. Area of anterior upper premolar when viewed from side
almost equaling that of posterior tooth. Second upper premolar with
main cusp straight, situated at or near middle of crown. First lower
premolar higher and broader, when viewed in profile, than second
and third, the third the smallest of the three. Second upper molar
largest, third upper molar with less than half crown area of second.
First lower molar smaller than second.
Measurements—Average and extreme measurements in milli-
meters of a series of eighteen females from the Colorado Desert are
as follows: total length, 97.3 (93.0-103.0) ; tail vertebrae, 37.0 (33.0—
41.0) ; tibia, 23.0 (20.8-24.2) ; foot, 15.6 (14.0-17.0) ; forearm, 49.9
(46.8-52.6) ; greatest length of skull 23.1 (22.5-23.7); zygomatic
breadth, 11.2 (10.8-11.6); mastoid breadth, 9.8 (9.2-10.0); inter-
orbital constriction, 3.5 (3.4-3.7); height of cranium at bullae, 9.4
(9.8-9.7). The only male at hand measures: total length, 95.0 milli-
meters; tail vertebrate, 35.0; tibia, 23.0; foot, 14.0; forearm, 48.4;
ereatest length of cranium, 23.0; zygomatic breadth, 11.1; mastoid
breadth, 10.0; interorbital constriction, 3.5; height of cranium at
bullae, 9.3.
Synonymy and History—This bat was described by Baird (1858,
p. 116) from a specimen taken at Fort Yuma (Imperial County, Cali-
fornia, opposite Yuma, Arizona), under the name Macrotus Califor-
Nicus.
Coues and Yarrow (1875, pp. 80-81), presuming the lighter color-
ation of M. californicus to be due to fading in alcohol, and overlooking
other differences, recorded the species occurring in California as
identical with M. waterhousii of the West Indies. Dobson (1878, p.
465) states that his own conclusion that waterhousii and californicus
are identical was based on descriptions alone, he not having seen the
type of californicus.
Teeth—Dental formula: j
ei) tee
1918] Grinnell: A Synopsis of the Bats of California 255
MEASUREMENTS IN MILLIMETERS OF EIGHTEEN FEMALES OF MAaAcrotTus
CALIFORNICUS BAIRD, FROM NEAR TORRES (= Toro),
RIVERSIDE COUNTY, CALIFORNIA
é E as
ae 2 2 5 Bs
1206 103.0 21.5 16.0 48.3 23.6 3
1207 97.0 22.0 16.0 50.0 23.6 11.3
1208 95.0 23.2 16.0 50.0 22.7 11.0
1209 100.0 23.6 17.0 52.6 23.0 11.0
1210 95.0 22.2 16.0 AOOy seis, eee) 0 SE ae
1211 99.0 22.2 16.0 49.1 22.6 11.8 9.8 3.6
1212 96.0 23.0 15.0 50.5 22.8 11.3 10.0 3.4
1213 100.0 24.2 17.0 50.0 23.2 11.0 9:9) 3.6
2M) AEE Anse a eee 15.0 50.2 23.2 10.8 10.0 3.5
1215 95.0 37.0 22.0 15.0 49.5 22.5 11.4 929, 3.7
1216 94.0 36.0 20.8 15.0 46.8 23.4 11.3 10.0 3.7
1217: 96.0 33.0 22.3 14.0 49.3 23.0)) ge 10.0 3.4
1218 99.0 38.0 21.7 16.0 50.3 23.3 11.2 9.8 3.4
1219 95.0 39.0 22.6 15.0 48.0 23.0 11.0 9.2 3.3
1220 95.0 39.0 23.6 16.0 52.0 23.0 11.2 10.0 3.7
1221 93.0 37.0 23.4 16.0 50.0 22.8 11.0 9.8 3.5
1222 101.0 41.0 23.7 16.0 51.0 23.6 11.4 10.0 3.5
1223 101.0 36.0 23.4 14.0 51.7 23.7 11.6 itl 3.7
Distribution.—The distribution of Macrotus californicus was given
by Rehn (1904, p. 441) as throughout the arid region of the south-
western United States, Lower California and Sonora. The species is
now recorded from as far eastward as Tombstone, Arizona (Rehn,
loc. cit.) ; westward to De Luz, San Diego County, California (see
below) ; south to Camoa, Rio Mayo, Sonora, and Cape San Lucas,
Lower California (Rehn, loc. cit.) ; north to Riverside, California
(Stephens, MS). In California the leaf-nosed bat seems to be con-
fined to the hottest parts of the Lower Sonoran zone, mainly on the
Colorado Desert. (See map, text-fig. H.)
Specimens Examined.—The writer has examined 113 specimens
from the following localities in California: San Diego County: Valle-
cito, 16 (U. S. Nation. Mus., 9; Mus. Vert. Zool., 7); De Luz, 10
(Stanford Univ., 8; U. S. Nation. Mus., 2); Santa Margarita Ranch
and River, 17 (Stanford Univ.) ; Imperial County: Indian Wells, 4
(U. S. Nation. Mus.) ; Mecca, 2; Palo Verde, 1; Riverside County:
Colorado desert near Torres [= Toro], 63.
Natural History —In September, 1893, Edward Hyatt (MS) cap-
tured eighteen specimens of Macrotus californicus in a cave on the
256 University of California Publications in Zoology [Vou.17
upper part of the Santa Margarita Ranch, Santa Margarita River,
San Diego County. The principal insect remains found scattered
about the floor of the cave were of the following beetles, all common
F = aw
2 yao
MACROTUS CALIFORNICUS
EUMOPS CALIFORNICUS
MYOTIS LUCIFUGUS ALTIPETENS
MYOTIS LUCIFUGUS ALASCENSIS
DISTRIBUTION MAP
| MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA
Fig. H. Map showing stations of occurrence in California of Macrotus cali-
fornicus, Eumops californicus, Myotis lucifugus altipetens, and Myotis lucifugus
alascensis, as established by specimens examined by the author.
species in southern California: Ligyrus gibbosus, Chlaenius sericeus,
Polyphylla decemilineata; also various species of flies.
Sixty-three specimens of this bat were secured by collectors from
the California Museum of Vertebrate Zoology on April 15, 1908, in
a cave in the Coachella Valley, near Toro, Riverside County (pl. 14,
1918] Grinnell: A Synopsis of the Bats of California 257
fig. 1). This valley is an old sea-bottom and at sea-level there are
several caves worn in the rocky hillside by the one-time action of
waves. Hight of these caves were examined by the party and five
showed evidence of having been occupied by bats. <A portion of the
floor of one of these caves was found to be covered to a depth of sev-
eral inches with bat guano, and this was the cave where all the
specimens were obtained. <A colony of fully three hundred of these
bats was clustered in a funnel-shaped recess in the roof about thirty
feet from the entrance. Disturbed by the light of the lantern the
bats took wing and streamed out. Of the sixty-three taken, all but
two were females and nearly every one contained a single large
embryo; none was found to contain more.
In the spring of 1914 the cave referred to above was visited by
Donald R. Dickey, who states in a letter that he found it untenanted ;
but a mile or so away another cave was found which contained a
colony of bats. A number of individuals were caught, examined,
and liberated. All proved to be California leaf-nosed bats.
On the morning of Mareh 29, 1908, C. H. Richardson (MS) found
a female leaf-nosed bat in a mouse trap set on the open ground near
Mecea, Imperial County. The bat must have been entrapped while
attempting to capture the ants or beetles which commonly visit traps
for the oatmeal with which they are baited. It would appear that
this species of bat may seek some of its food upon the ground.
Dobson (1878, p. 466) states that a correspondent writing from
St. Ann’s, Jamaica, remarks that Macrotus waterhousti there eats not
only insects but also fruits. He mentions particularly the fustiec
berry (Morus tinctoria), the bread-nut (Brosimum alicastrum), and
the rose-apple (Hugenia jambos). It would be very interesting to
learn whether our Californian Macrotus also eats fruit as well as
insects ; if so, it is the only fruit-eating bat occurring within the state.
Stephens (1906, p. 277) considers these bats as being probably
migratory; for he failed to find them in January in a place where
they were nearly always to be found in spring and summer. ‘The
writer knows of no dates of capture within the state later than Sep-
tember or earlier than March, save those furnished by Leo Wiley.
There is in the Museum of Vertebrate Zoology an alcoholic. speci-
men of Macrotus californicus (no. 23651), taken by this observer
December 31, 1915, in an old mine shaft near Palo Verde. Mr.
Wiley writes (MS) that he secured five more individuals of this
species in the same shaft on February 20.
258 University of California Publications in Zoology [Vou.17
Tn regard to the usual number of young, Stephens (1906, p. 277)
says: ‘‘The young are born in June. More than half of the females
bear two young, the remainder but one.’’ In the ease of the above
described colony near Toro all the pregnant females examined con-
tained but one foetus each.
Family VESPERTILIONIDAE
The distribution of this family is the same as that of the order
Chiroptera.
The members of this family include bats having muzzles without
distinct leaflike outgrowths and having ears with well developed
anterior basal lobes; the tragus is simple and usually well developed ;
the tail is well developed, but extends only to, or but very slightly
beyond, the edge of the wide interfemoral membrane.
Miller (1907, p. 196) divides the family into six subfamilies, rep-
resented by forty genera. The two subfamilies occurring in America
north of southern Mexico are the Vespertilioninae and the Nyeto-
philinae, and both are represented in California.
Subfamily VESPERTILIONINAE
Distribution same as for the Family. Primitive in structural
features, being characterized chiefly by lack of specialized structures
distinguishing the other subfamilies. Differs from the Nyetophilinae
in the absence, in all of its members, of abrupt truncation of the
muzzle, and of any ridge above the nostrils.
Genus Myotis Kaup
Representative species of this genus are found in the temperate
and tropical regions of both hemispheres; the area of their distribution
is probably not exceeded by that of any other genus of bats. About
eighty species and subspecies of Myotis are now known, fifteen of
which oceur in California.
22 flak pi 3-3* moe 2 oe
Say) dil? sea) a3)
Upper incisors well developed, subequal, and closely crowded ; crowns
almost cylindrical in cross-section; inner incisor with a distinet
posterior secondary cusp, the outer with a well developed concave sur-
face directed toward canine, from which it is separated by a space not
Characters —Dental formula:
*In at least one species, Myotis occultus, the upper middle premolar, pm*, is
sometimes wanting.
1918] Grinnell: A Synopsis of the Bats of California 259
quite equal to the combined diameters of both incisors. Lower incisors
forming a continuous row strongly bowed forward between canines;
first and second lower incisors with rather narrow, trilobed cutting
edges, the third much wider than the others and with three or four
tubercles. Canines, both above and below, without secondary cusps,
but with small yet distinct cingula. The two anterior upper pre-
molars small, especially the central one, which is often minute and
crowded out of the tooth-row internally; lower premolars following
relative proportions of three upper ones, but central one, although
smallest, rarely so minute as corresponding tooth in upper jaw. Last
upper molar rather less in area of cross-section than half the second.
The skull varies considerably in size among the different species,
but usually has brain-case rounded and elevated above facial region,
which is narrow, depressed and markedly saddle-shaped; depth of
brain-ease, including auditory bullae, about equal to mastoid breadth ;
sagittal erest low but usually distinct. Auditory bullae well devel-
oped and covering more than half the surface of the cochleae, but
simple in form and not very large; diameter of bullae about equal to
distance between them. Face always hairy; glandular prominences
much less developed than in related genera and seareely adding to
breadth of face. Apertures of nostrils, crescent-shaped. Ear oval,
distinctly higher than broad. Tragus slender, and nearly or quite
straight. Foot, lower leg, and calear of variable size and develop-
ment. Wing broad, the fifth metacarpal being slightly longer than,
or equal to, third, the posterior attachment anywhere from a little
above ankle to base of toes. Tail long, usually exceeding forearm in
length.
Myotis velifer (J. A. Allen)
Cave Bat
Vespertilio velifer J. A. Allen (1890, pp. 177-178). Original description ;
type locality, Guadalajara, Jalisco, Mexico.
Myotis velifer, J. Grinnell (1913b, p. 276). Range in California.
Myotis velifer, J. Grinnell (1914, p. 266). Occurrence at Needles, Cali-
fornia. :
Diagnosis —Aceording to Miller (1897), p. 57), Myotis velvfer
is the largest species of Myotis known to occur anywhere in the United
States or Mexico. Total leneth, 90-105 millimeters; length of fore-
arm, 40-47 millimeters. Calear slender, without a well developed lobe.
Free border of uropatagium naked. Kars short, reaching only to tip
of nose. Wing membrane arising from metatarsus.
Description: Head.—Kars short (about 16 millimeters in height)
and pointed. Tragus about half height of ear, its anterior edge
straight or very slightly concave towards the tip.
Limbs and membranes.—Wing attached nearer to bases of toes
than to ankle. Free border of interfemoral membrane shorter than
ealear, and naked.
Pelage.—Ears scantily haired over whole ventral surface, as also
at extreme bases on dorsal surface. Wings haired on ventral surfaces
at their extreme bases; otherwise naked. Uropatagium sparsely
260 University of California Publications in Zoology (Vou. 17
haired, both dorsally and ventrally, on its proximal fourth. Toes
sprinkled on their dorsal surfaces with stiff hairs from 2 to 3 milli-
meters long.
Color.—Miller (1897b, p. 57) gives the normal coloration of this
bat as ‘‘dull sepia throughout, paler on the belly, the hairs everywhere
dusky slate at base.’’ One specimen at hand from California has
the hairs everywhere chaetura black at their bases. On the dorsal
surface the distal portions of the hairs are drab, while on the ventral
surface the distal portions are cartridge buff. On another skin from
California the distal portions of the hairs upon the dorsal surface are
avellaneous.
Skull.—vVery large and strong for a Myotis (see table of measure-
ments) ; sagittal crest unusually well defined (pl. 21, fig. 32). For
comparisons with M. thysanodes and M. evotis, the only other species
with skulls of similar length, see pages 294 and 298.
Measurements.—Average and extreme measurements of the three
specimens in the Museum of Vertebrate Zoology are as follows: total
length, 96.6 (93.0-100.0) ; tail vertebrae, 40.6 (40.0—42.0) ; tibia, 15.6
(15.0-16.0) ; foot, 10.0 (9.0-10.5) ; forearm, 40.4 (39.6-41.0) ; greatest
length of cranium, 16.3 (16.0-16.6).
MEASUREMENTS IN MILLIMETERS OF THREE EXAMPLES OF MYOTIS VELIFER
(J. A, ALLEN), FROM NEEDLES, SAN BERNARDINO COUNTY, CALIFORNIA
: “2
Fy 5 crates oe
= a B Ss Sie
a= 2 E Ee : fs 6S
M £¢ mo 5 =) y o R oe
aa Sex ae ie aq im = pq
7762 «of 100.0 42 15.0 10.5 40.8 10.3 8.1
7763 og 93.0 40.0 16.0 9.0 41.0 10.0 8.1
7764 Q 97.0 40.0 15.9 10.5 39.6 10.4 8.3
Synonymy and History.—The cave bat was described by J. A.
Allen (1890, pp. 177-178) under the name Vespertilio velifer, from
three specimens and six additional skulls obtained at Santa Cruz del
Valle, Guadalajara, Jalisco, Mexico. H. Allen (1894, p. 92) placed
velifer as a subspecies of Vespertilio albescens of South America.
Miller (1897), p. 59) thinks it best to retain the specific name velifer
for the bat occurring in Mexico and the United States until the South
American species has been definitely characterized.
Distribution—The general range of this bat as given by Miller
(1912, p. 54) is from Hidalgo, northern Michoacan, and the City of
Mexico north to the southern border of the United States. In Cali-
fornia it has been taken only at Needles, San Bernardino County, in
the Lower Sonoran zone near the Colorado River. (See map, text-
fig. N.)
Natural History.—Bailey (1905, pp. 208-209) remarks that if
this bat is habitually a ecave-dweller, the distribution of eaves probably
accounts for its somewhat erratic range.
1918] Grinnell: A Synopsis of the Bats of California 261
Ward (1891, p. 744), in his report of a collecting trip made to
Las Vegas, Canton of Jalapa, Vera Cruz, Mexico, in February, 1891,
describes a visit made to certain tunnel-like caves on the side of a
long extinct voleano. In the cool atmosphere of these caves, fresh-
ened by the current of air which continually flowed down the
mountain side, bats were found in a state of semi-torpor. The pre-
vailing species was Myotis velifer, which was present in hundreds
Ward secured one hundred and fifty-two males and forty-two females.
The author adds: “‘This I take to be a fair average of the proportion
of the sexes in what is probably one of their permanent headquarters.’
The three specimens of the cave bat in the Museum of Vertebrate:
Zoology were taken by C. L. Camp at Needles, California, on July 16:
and 18, 1909. Bats of this species were found roosting in numbers
in an old storehouse there.
Myotis occultus Hollister
Holiister Bat
Myotis oceultus Hollister (1909, p. 43). Original description; type locality,
Colorado River, ten miles above Needles, San Bernardino County, Cali-
fornia.
Myotis occultus, Miller (1912, p. 55). Nominal.
Myotis occultus, J. Grinnell (1913b, p. 276). Range in California.
Myotis occultus, J. Grinnell (1914, pp. 263-265, figs. H, I). Occurrence
on the Colorado River.
Diagnosis —Total length 85 to 90 millimeters, forearm 35.1 to
36.4; calear longer than free border of interfemoral membrane; wing
membrane arising from near bases of phalanges; cranium relatively
flat-topped and rostrum broad.
Description: Head.—FKorm externally not in any way peculiar.
In a series of six (dried) specimens the average height of the ear is
11.5 millimeters; of the tragus, 6.1 (both measured from notch).
Limbs and Membranes——Wing membrane arises from near bases
of toes. Calear longer than free border of interfemoral membrane.
Pelage—F ur everywhere full and soft. On the body the majority
of the hairs are about 3 millimeters long, but scattered among these
shorter hairs are occasional longer ones, averaging about 5 millimeters
in length. Wings furred only at their extreme bases, both above and
below. Uropatagium seantily haired on its basal fifth, the hairs
being longer on the dorsal surface than on the ventral. Ear haired
on posterior surface at extreme base; a line of hairs extends along
basal third of anterior border of ear. Entire anterior surface of ear
sparingly haired. Edge of upper lip furnished with a scanty row
of downward projecting hairs, and a few longer hairs, 4 to 8 milli-
meters in length, project from glandular portion of face.
262 University of California Publications in Zoology [Vou.17
Color—tThe six specimens of the Hollister bat which are at hand
are quite uniform in coloration. Bases of hairs everywhere chaetura
black, except on anterior portion of throat where they are uniform
cinnamon buff. Distal portions of hairs on dorsal surface of body
light sayal brown, with a silky sheen. This shade of brown extends
down onto sides of belly, which are otherwise pale cinnamon buff,
varying slightly in exact shade among the different specimens. Mem-
branes clove brown.
Skull—asily distinguished from that of any other species of
Myotis inhabiting California, by its very broad and relatively flat-
topped rostrum and brain-case (pl. 21, fig. 30; pl. 22, fig. 42). Sag-
gital crest more clearly defined than in any other Californian Myotis,
except M. velifer. Teeth essentially as described for the genus, save
that in four of the eight known SUES HLEDE the upper middle premolar
(pm2) is missing on both sides.
Measurements —T he average and extreme measurements in milli-
meters of the six examples of the Hollister bat in the Museum of
Vertebrate Zoology are as follows: total length, 87.3 (85.0-90.0) ; tail
vertebrae, 35.6 (34.0-38.0) ; tibia, 13.9 (13.7-14.0) ; foot, 8.7 (8.0-
9.0); forearm, 35.6 (35.1-36.4); greatest length of cranium, 15.1
(14.8-15.4).
MEASUREMENTS IN MILLIMETERS OF Stx EXAMPLES OF MYOTIS ocCULTUS
HOLLISTER, FROM IMPERIAL COUNTY, CALIFORNIA
E - “2 ae
= ee og SF
= 3 = a 3s E=iin4 Lan;
ae 2 = 2 He <5 Su
Mus. fe 5 Es = i z S SE ee $8
no. Sex mien cue = oF & te Si [a2] fap
10702 @ 85.0 35.0 14.0 9.0 35.4 15.2 9.8 7.7 3.9
10703" @ 89.0 34.0 13.8 8.9 3:01 eee 9.7 -_ 4.1
10704" 90.0 36.0 14.2 9.0 36.4 15.4 10.1 7.9 4.0
10705° Q 86.0 34.0 13.9 8.5 35.2 14.8 9.6 7.3 4.1
10706 87.0 37.0 13:7, 8.0 36.3 15.3 9.6 7.6 4.0
107077 @ 87.0 38.0 14.0 9.0 35.1 15.0 9.8 ath 4.0
1From four miles south of Potholes.
? From five miles northeast of Yuma.
Synonymy and History—The Hollister bat was deseribed by
Hollister (1909, p. 48) under the name JJyotis occultus, from two
specimens obtained on the west side of the Colorado River, ten miles
above Needles, San Bernardino County, California, May 14, 1905.
Distribution—Known only from the west side of the Colorado
River from near Needles (as above) to near Yuma, where six speci-
mens were obtained in May, 1910 (see J. Grinnell, 1914, p. 263). (See
map, text-fig. O.)
Specimens Examined.—tThe writer has examined eight specimens
of the Hollister bat, from the following localities in California: Im-
perial County: five miles northeast of Yuma, 5; four miles south of
1918] Grinnell: A Synopsis of the Bats of California 263
Potholes, 1; San Bernardino County: west side of Colorado River
above Needles, 2 (U.S. Biol. Surv.).
Natural History—J. Grinnell (loc. cit.) suggests that the Hollister
bat is a late spring arrival in the region where it was found, since Hol-
lister’s two specimens were taken in May, and the Museum expedition,
although collecting along the Colorado River from February 15 until
May 15, failed to detect this bat until the first week of May. This
author states that of the six specimens obtained ‘‘the first was shot at
late dusk close to the river bank between files of cottonwoods, in
just the same association as those taken by Hollister. At our second
locality of capture, the remaining five specimens were shot over the
water in a back eddy of the river. Here these bats arrived in con-
siderable numbers at early dusk to drink, flitting down to the water’s
surface and dipping several times before flying off among the willows
and cottonwoods. ”’
Myotis lucifugus altipetens H. W. Grinnell
High Sierra Bat
Myotis ywmanensis satwratus, C. H. Merriam (1899, p. 89). Occurrence on
Mt. Shasta.
Myotis ywmanensis, Rehn (in Stone, 1904b, p. 590). Record of specimen
from Mt. Tallac.
Myotis yumanensis saturatus, Stephens (1906, p. 267), part. Diagnosis;
distribution.
Myotis yumanensis saturatus, J. Grinnell (1913b, p. 277), part. Range in
California.
Myotis yumanensis altipetens H. W. Grinnell (1916, pp. 9-10). Original
description. Type from Merced Lake, Yosemite National Park, Cali-
fornia.
Diagnosis —Length, 79 to 91 millimeters; tibia short, 15.0 to 16.4;
distal portion of fur on back isabella color, on belly pale ght buff.
Description: Head.—Ear short (about 12 millimeters in height),
and bluntly rounded at tip (pl. 18, fig. 15) ; anterior border straight
through lower third, then gently convex to the tip; posterior border
shghtly concave along upper two-thirds, then convex to basal notch.
In a dried specimen at hand the tragus is 6 millimeters in height and
slender, varying in width from 1.5 millimeters near the base to 0.5
millimeters just below the rounded tip. Nostrils as in other Myotis.
Limbs and Membrancs.—Membranes leathery but not thick. Cal-
ear slender and indistinct, greater in length than free border of
uropatagium; keel not prominent, and terminating lobule scarcely
apparent. Feet large (9 to 12 millimeters in length) and strong,
and more than half the tibia in length. The wing arises from a point
264 University of California Publications in Zoology [Vow.17
nearer to bases of toes than to ankle. Claws on both feet and thumbs
long, well curved, and of needle-like slenderness.
Pelage-—F ur everywhere full and soft. Longer hairs on middle
of back about 7 millimeters in length. Ear haired dorsally only at
base, but ventrally short fine hairs scattered sparsely over entire
surface. Wing membranes naked save where body fur extends alone
line of attachment of wings to body. Uropatagium very scantily
haired on its ventral surface. Toes scantily haired, both above and
beneath, on terminal joints.
Color—Distal portion of fur on back isabella color; bases of the
hairs chaetura black. Ventrally, the bases of the hairs are chaetura
black and the tips are a pale tint of light buff. Wings, feet, and
membranes deep clove brown, which becomes lighter at posterior edges
of wing membranes and uropatagium.
Skull—Of medium size for the genus (length 14.2 to 15.0 milli-
meters). Brain-ecase and rostrum broad and heavily built. Rostrum
not sharply turned up at tip, and forehead sloping back very gently.
Sagittal crest but faintly indicated, and occipital elevation very slight.
The brain-ease lacks the inflation at the front which is a character of
some other members of the genus. (See pl. 21, fig. 31; pl. 22, fig. 43.)
Measurements.—Average and extreme measurements of a series of
eight adult examples of Myotis lucifugus altipetens from California
are as follows: Total length, 86.8 (79.0—-91.0) ; tail vertebrae, 35.6
(31.0-41.0) ; tibia, 15.6 (15.3-16.4) ; foot, 10.4 (9.0-12.0) ; forearm,
35.8 (35.2-36.5) ; greatest length of cranium, 14.5 (14.2-15.0).
Synonymy and History.—This species was described under the
name Myotis yumanensis altipetens, by H. W. Grinnell (1916, pp.
9-10). The type specimen was taken one mile east of Merced Lake,
Yosemite National Park, at an altitude of 7500 feet. Four of the
five specimens of bats secured by C. H. Merriam (1899, p. 89) in the
Hudsonian Zone on Mount Shasta and by him recorded as Myotis
yumanensis saturatus have been examined by the writer and prove
to be MW. 1. altipetens, as does the specimen recorded by Rehn (in Stone,
19046, p. 590) under the name Myotis yumanensis, from Mount
Tallae. F
Distribution.—As indicated by the material at hand, the High
Sierra bat inhabits the central Sierra Nevada, the vicinity of Mount
Shasta, and the Warner Mountains. It oceurs chiefly in the Can-
adian and Hudsonian zones, and has been secured at a greater altitude
than any other bat in the collection of the Museum of Vertebrate
Zoology. Merriam (1899, p. 89) says of his Mount Shasta specimens:
‘“‘The species is interesting as the only bat secured in the Hudsonian
zone.’’ The following list of localities shows the record stations thus
far established in California. (See map, text-fig. H.)
f the Bats of California
: A Synopsis o
Grinnell
1918]
MEASUREMENTS IN MILLIMETERS or E1cut EXAMPLES OF MYOTIS LUCIFUGUS ALTIPETENS H. W.
* Mus.
13805
23035
23034"
12036
23036
13804
1 Type.
3 0 Q A A QA A AZ
Locality
E. face Warren Peak, 8700 ft. alt.,
Warner Mts., Modoe. Co.
E. face Warren Peak, 8700 ft. alt.,
Jarner Mts., Modoe Co.
Castle Lake, 5434 ft. alt., Sis-
kiyou Co.
Vogelsang Lake, 10350 ft. alt.,
Mariposa Co.
1 mi. E. Merced Lake, 7500 ft. alt.,
Mariposa Co.
Gilmore Lake, 8400 ft. alt., Eldo-
rado Co.
Vogelsang Lake, 10350 ft. alt.,
Mariposa Co.
Castle Lake, 5434 ft. ‘alt., Sis-
kiyou Co.
lengta
Total
Foot
Ear from
crown
S
B
®
Ki
5
&
length of
cranium
Greatest
(=
Le
ea
14.7
Zygomatic
breadth
oo
©
Breadth of
brain-case
GRINNELL, FROM CALIFORNIA
Interorbital
constriction
266 University of California Publications in Zoology [Vou.17
Specimens Examined.—The writer has examined 23 specimens
from the following localities in California: Modoe County: east face
of Warren Peak, 2; Warner Mts., 1 (Calif. Acad. Sci.) ; Siskiyou
County: Castle Lake, 2; Mt. Shasta, 8 (U. S. Biol. Surv., 4; Mus.
Vert. Zool., 4); Nevada County: Independence Lake, 2; Eldorado
County: Gilmore Lake, 2; Mt. Tallac, 2 (Mus. Comp. Zool., 1; Acad.
Nat. Sei. Phila., 1); Mariposa County: Merced Lake, 2; Vogelsang
Lake, 2.
Comparisons.—M yotis lucifugus altipetens may be distinguished
from Myotis lucifugus alascensis by the difference in depth of color;
the shade is mummy brown above in alascensis, isabella color in alti-
petens. The race altipetens is somewhat intermediate in color be-
tween the race carissima of the Rocky Mountain region, and alascensis
of the Northwest. Bats of the lucifugus group may always be dis-
tinguished externally from those of the longicrus group by the much
longer tibia of the latter (15.0-16.4 in lucifugus, 17.9-18.9 im longi-
crus). The skull of longicrus has a much shorter, more sharply
turned-up rostrum than that of lucifwgus, and the facial angle is
greater (see pl. 22). The separation between the lucifugus and
yumanensis groups is not wide, and identification of young individuals
is not possible without the aid of cleaned skulls. Externally, the
claws on the foot and thumb are often an aid to identification, being
in lucifugus much longer, more slender, and more curved. The skull
of yumanensis is slightly smaller and uniformly more slender than
that of lucifugus, with the brain-case more inflated in the frontal
region.
Natural History.—C. H. Merriam (1899, p. 89) says of the oceur-
rence of the High Sierra bat on Mount Shasta: ‘‘Common among
the alpine hemlocks at Squaw Creek Camp, where they were seen
every night, darting in and out of the flickering light of the camp
fire.’ When camped at Merced Lake and at Vogelsang Lake, in
the Yosemite National Park, J. Grinnell and W. P. Taylor secured
several of these bats. The former states (MS) that one of the speci-
mens taken was shot at 7:10 on the evening of August 19. This bat,
which proved to be a male, was flying high, straight down the canyon
between the lodgepole pine tops, and was being pursued by another
bat, apparently of the same species. A specimen secured at Vogel-
sang Lake, altitude 10,350 feet, September 2, was shot at 7:11 pP.m.,
as it was skimming over the surface of the water.
1918] Grinnell: A Synopsis of the Bats of California 267
Myotis lucifugus alascensis Miller
Alaska Brown Bat
Myotis lucifugus alascensis Miller (1897b, pp. 63-64). Original deserip-
tion; type from Sitka, Alaska.
Diagnosis —Similar to Myotis lucifugus altipetens, but color
darker; general color of fur on back mummy brown.
Description.—Feet, ears, and membranes chaetura black, as also
the bases of the hairs everywhere. Terminal portion of fur on back
mummy brown; on ventral surface tips of hairs drab. Fur on middle
of back 6 to 7 millimeters in length. (For comparisons between this
and other nearly related races see under Myotis lucifugus altipetens,
p. 266.)
Measurements—The single example at hand from California
measures in millimeters as follows: total length, 86.0; tail vertebrate,
39.0; foot, 9.0; forearm, 35.0; tibia, 15.0; greatest length of skull, 14.9.
Synonymy and History—The Alaska brown bat was deseribed by
Miller (1897), pp. 63-64), who examined sixteen speeimens, thirteen
being from Alaska, and the remaining three from Queen Charlotte
Islands, British Columbia. The type specimen was procured at
Sitka, Alaska.
Distribution —Miller (1912, p. 55) gives the range of Myotis luci-
fugus alascensis as the humid coast district of southern Alaska and
northern British Columbia. The specimen now at hand from Cali-
fornia is an adult male (no. 11843, Mus. Vert. Zool.), taken at Eureka,
Humboldt County, August 2, 1910 (see map, text-fig H). I can
find no previous record of this race from California.
Natural History.—Nothing is known of the habits of this bat in
California.
Myotis longicrus longicrus (True)
Northwestern Long-legged Bat
V [espertilio]. lucifugus, Cooper (in Cronise, 1868, p. 442)(?). Occurrence
in California.
Vespertilio lucifugus, Cooper (1868, p. 5)(?). Fort Reading (Shasta
County).
Vespertilio subulatus, Cooper (in Cronise, 1868, p. 442)(?). Listed as
occurring in California.
Vespertilio longicrus True (1886, p. 588). Original description; type
locality Puget Sound.
Vespertilio lucifugus, C. H. Townsend (1887, p. 182). Reeord of occur-
rence at Fort Reading.
Myotis lucifugus longicrus, Miller (1897b, pp. 64-65), part. Description;
distribution. Specimens listed from Nicasio, Point Reyes, ete.
268 University of California Publications in Zoology [Vov.17
Myotis lucifugus longicrus, Merriam (1899, p. 89). Oceurrence on Mt.
Shasta.
Myotis lucifugus longicrus, Elliot (1901, pp. 402-403), part. Diagnosis;
distribution.
Myotis lucifugus longicrus, Stone (1904a, p. 579). Occurrence on Mt.
Sanhedrin.
Myotis lucifugus longicrus, Elliot (1904b, p. 581), part. Diagnosis; dis-
tribution.
Myotis lucifugus longicrus, Elliot (1905, p. 479), part. Geographic dis-
tribution. +
Myotis lucifugus longicrus, Stephens (1906, pp. 265-266), part. Diag-
nosis; distribution.
Myotis lucifugus longicrus, Seton (1909, p. 1149), part. Map showing
general range.
Myotis lucifugus longicrus, Miller (1912, p. 55), part. General range.
Myotis lucifugus longicrus, J. Grinnell (1913b, p. 276), part. Range in
California.
Myotis longicrus, Miller (1914, pp. 211-212). Possible identity of M. 1.
longicrus from interior of California with M. 1. interior.
Diagnosis.—Total length, 91 to 100 millimeters; tibia long, 17.4
to 18.9; ear from meatus, 12.0 to 13.5; tragus, 7.0 to 8.0; general color
of back deep bister.
Description: Head.—Kar short, reaching barely to tip of nose
when laid forward, and bluntly rounded at tip. Tragus bluntly
rounded at tip, and its greatest diameter about 1 millimeter above its
basal notch.
Limbs and Membranes.—Membranes not peculiar. Feet large and
strong (7 to 8 millimeters long), but less than half tibia in length.
Wing membrane attached at bases of toes. Calear keeled.
Pelage-——F ur everywhere full and soft; length on middle of back
averaging about 7 millimeters. Wing membranes naked save where
body fur extends along line of attachment of wings to body. Dorsal
surface of uropatagium scantily haired on its basal fourth; ventrally,
short, scanty hairs occur over nearly its entire surface. Terminal
joint of each toe scantily haired, both above and below. Ear scantily
haired over entire anterior surface; posterior surface haired at base,
the hairs extending farther outward along anterior margin than on
posterior edge.
Color—Hairs on back deep bister, with faintly hghter tippings.
Hairs on ventral surface deep bister at bases, with tippmgs of
vinaceous-buff, these light tippings being more conspicuous toward
posterior portion of ventral surface. Ears and membranes dark clove
brown. Individuals from the Sierra Nevada of Placer and Eldorado
counties are faintly lighter than those from the humid coast belt, and
thus vary in the direction of MW. 1. interior of the more arid southern
and interior parts of the state.
Skull.—Distinguished from that of any other small Myotis by its
relatively short and sharply upturned rostrum and very high occipital
elevation (pl. 22, fig. 40). Between skulls of the nearly related
Myotis longicrus longicrus and M. l. interior I ean find no constant
difference; but Miller (1914, p. 211) says of the latter race, ‘‘skull
1918 | Grinnell: A Synopsis of the Bats of California 269
tending to be slightly larger than in true longicrus and with less
abruptly rising frontal and occipital regions; teeth normal.’’
Measurements.—Average and extreme measurements in milli-
meters of a series of six adults from California are as follows: Total
length, 95.0 (87.0-100.0) ; tail vertebrae, 43.0 (37.0-47.0) ; tibia, 17.9
(17.4-18.9) ; foot, 7.9 (7.5-8.5) ; forearm, 37.2 (35.7-39.2) ; greatest
leneth of cranium, 13.9 (13.7—-14.2).
MEASUREMENTS IN MILLIMETERS OF STIX ADULTS OF MYOTIS LONGICRUS LONGICRUS
(TRUB) FROM CALIFORNIA
™ =a Aa oad 2
a 2 E s
20811" io] 91.0 AZOU ances (Sie eres
69497 2 100. 47.0 18.5 8.0 39.5
18679° ie) 99.0 46.0 17.4 8.5 37.5
24185* 2 98.0 45.0 18.9 8.0 37.6
20810! 3 95.0 41.0 17.4 Ted) 35.7
19699° fof 87.0 37.0 17.4 35.8
1From Guerneville, Sonoma County.
*From Pescadero Creek, San Mateo County.
*From Dutch Flat, Placer County.
4From Fyffe, El Dorado County.
* From Cazadero, Sonoma County.
Synonymy and History—The northwestern long-legged bat was
described by True (1886, p. 588) under the name Vespertilio longi-
crus, from a specimen obtained at Puget Sound.
Distribution.—The California range of Myotis longicrus longicrus
cannot be confidently stated without much further field work. As
will be noted from the list of verified localities, this bat appears
to be restricted to the Transition and Upper Sonoran zones in the
humid coast belt south as far as Pescadero Creek, San Mateo County,
and in the northern portion of the Sierra Nevada, south at least to
Walker Lake, Mono County (see map, text-fig. I). The range of
Myotis ‘‘lucifugus’’ longicrus as given by Miller (1912, p. 55) and
J. Grinnell (1913), p. 276) includes that of the newly recognized
Myotis longicrus interior.
Specimens Examined—The writer has examined 14 specimens
from the following loealities in California: Mendocino County: Mt.
Sanhedrin, 1 (Acad. Nat. Sci. Phila.) ; Sherwood, 2; Sonoma County :
near Guerneville, 2; near Cazadero, 1; Nevada County : Independence
Lake, 1; Placer County: Dutch Flat, 1; Eldorado County: Fyffe,
3; Mono County: Walker Lake, 1; San Mateo County: Menlo Park, 1;
Pescadero Creek, 1.
270 University of California Publications in Zoology [Vou.17
Natural History—Bats, like birds, have their chosen feeding
ranges and the range of one species supplements that of another
instead of duplicating it. When encamped near the Russian River
in Sonoma County in 1913, I had opportunity to study the feeding
@ MYOTIS LONGICRUS LONGICRUS
4 MYOTIS LONGICRUS INTERIOR
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA
qe ee
Fig. I. Map showing stations of occurrence in California of Myotis longicrus
longicrus and Myotis longicrus interior, as established by specimens examined by
the author.
habits of four species of bats, among them Myotis longicrus longicrus.
This bat first appeared at 7:45 p.m. (June 11), twenty minutes after
sunset, and none was seen after 8:30- p.m. Individuals flew in and
out among the tents and young redwoods of the resort at a height of
1918] Grinnell: A Synopsis of the Bats of California 271
from six to twenty-five feet above the ground. One fly-way, which
was used every evening, I was inclined to believe was the route of a
single individual.
Myotis longicrus interior Miller
Interior Long-legged Bat
Myotis lucifugus longicrus, Miller (1897b, pp. 64-65), part. Description;
distribution; specimens listed from San Emigdio, Owens Lake, ete.
Myotis lucifugus longicrus, Elliot (1901, pp. 402-403), part. Diagnosis;
distribution.
Myotis lucifugus longicrus, Elliot (1904a, p. 318). Occurrence in Inyo
Mountains.
Myotis lucifugus longicrus, Elliot (1904b, p. 581), part. Diagnosis; dis-
tribution.
Myotis lucifugus longicrus, Elliot (1905, p. 479), part. Geographic dis-
tribution.
Myotis lucifugus longicrus, Elliot (1907, pp. 504-505). Repeated record.
Myotis lucifugus longicrus, Seton (1909, p. 1149), part. Map showing
general range.
Myotis lucifugus longicrus, J. Grinnell (1908, p. 158), part. Occurrence
in the San Bernardino Mountains.
Myotis lucifugus longicrus, Miller (1912, p. 55), part. General range.
Myotis lucifugus longicrus, Grinnell and Swarth (1913, p. 380). Oceur-
rence in the San Jacinto Mountains.
Myotis lucifugus longicrus, J. Grinnell (1913b, p. 276), part. Range in
California.
Myotis longicrus interior Miller (1914, pp. 211-212). Original description ;
type from Twining, Taos County, New Mexico.
Diagnosis —Sinilar to Myotis longicrus longicrus, but color tawny-
olive, instead of deep bister.
Description.—Bases of hairs everywhere deep bister; distal half of
fur on back tawny-olive; on the ventral surface the tippings are of
cartridge buff, with the darker bases of the hairs showing through.
Ears and membranes clove brown, of a somewhat lighter tint than
those of longicrus.
Measurements.—Average and extreme measurements in milli-
meters of a series of twelve examples from California are as follows:
Total length, 98.9 (94.0-103.0) ; tail vertebrae, 45.7 (42.0-49.0) ; tibia,
18.1 (16.4-19.0) ; foot, 7.7 (7.0-8.0) ; forearm, 38.0 (36.8-39.3) ; great-
est leneth of cranium, 13.9 (13.3-14.4).
Synonymy and History.—The interior long-legged bat was de-
scribed by Miller (1914, pp. 211-212) from a specimen collected at
Twining, Taos County, New Mexico.
Distribution—Miller (loc. cit.) lists specimens of Myotis longi-
crus interior from Nevada, Idaho, Wyoming, Colorado, New Mexico,
Arizona, and Chihuahua. The exact range of this bat in California
[ Vou. 17
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1918 | Grinnell: A Synopsis of the Bats of California 273
cannot at present be given. It has been collected in the arid Upper
Sonoran, Transition, and lower Canadian zones, from Benton, Mono
County, south to the Cuyamaca Mountains, in San Diego County.
(See map, text-fig. I.)
Specimens Examined.—The writer has examined 23 specimens
from the following localities in California: Mono County: Benton, 1;
Inyo County: Little Onion Valley, 3; Kearsarge Pass, 1; Tulare
County: Monache Meadows, 1; Kern County: San Emigdio, 4; Los
Angeles County: Pasadena, 2; San Bernardino County: San Bernar-
dino Mts., 4 (Mus. Vert. Zool., 3; Acad. Nat. Sei. Phila., 1) ; Riverside
County: Santa Rosa Mts., 3; base of San Jacinto Mts., near Cabe-
zon, 1; San Diego County: Cuyamaea Mts., 3.
Natural History.—I ean find nothing recorded concerning the
natural history of this bat.
Myotis yumanensis yumanensis (H. Allen)
Yuma Bat
Vespertilio ywmanensis H. Allen (1864, pp. 58-59, figs. 54-56). Original
description; type loeality, Fort Yuma, Imperial County, California.
Vespertilio Ywmanensis, Cooper (1868, p. 5). Distribution.
V [espertilio]. macropus, Cooper (in Cronise, 1868, p. 442). Listed as oceur-
ring in California.
V [espertilio|]. Ywmanensis, Cooper (in Cronise, 1868, p. 442). Listed as
oceurring in California.
Myotis yuwmanensis, Miller (1897b, pp. 66-68), part. Description; dis-
tribution; recorded from Keeler, Lone Pine, Owens Lake, ete.
Myotis yumanensis, Elliot (1901, p. 403), part. Diagnosis; distribution.
Myotis ywmanensis, Miller and Rehn (1901, p. 256). Type locality.
Myotis yumanensis, Elliot (1904a, pp. 318-319), part. Occurrence in
Argus Mountains.
Myotis yumanensis, Elliot (1904b, pp. 576-577), part. Diagnosis; dis-
tribution.
Myotis yumanensis, Elliot (1905, p. 475), part. Distribution.
Myotis ywmanensis, Stephens (1906, p. 267), part. Diagnosis; distri-
bution.
Myotis yumanensis, Elliot (1907, pp. 501-502), part. Localities of capture
in California.
Myotis yuwmanensis, Lyon and Osgood (1909, p. 291). Record of type.
Myotis yuwmanensis yumanensis, Miller (1912, p. 56), part. Range.
Myotis yumanensis yumanensis, J. Grinnell (1913b, pp. 276-277), part.
Range in California.
Diagnosis —Size rather small (total length 74 to 88 millimeters) ;
calear distinet, considerably longer than free border of interfemoral
membrane, and terminating in a well-marked lobule. Feet very
274 University of California Publications in Zoology [Vou.17
large (7.9 to 9.1 millimeters long) and strong, larger in proportion
to size of body than in any other member of the genus Myotis. Gen-
eral color of back cartridge buff; of lower surface, whitish.
Description: Head.—EKar when laid forward reaching just beyond
tip of nose; tip of ear narrow and abruptly rounded off. Tragus
slender, about half height of ear, and pointed, its posterior border
erenulate.
Limbs and Membranes.—Membranes thicker than those of other
small members of the genus. Foot large, broad, and strong, its length
more than half that of tibia.
Pelage.—Hairs longest on middle of back, where they average
about 5 millimeters in length.
Color—An example (no. 171508, U. S. Nation. Mus.) taken at
Fort Mohave, Arizona, March 14, 1911, is colored as follows: hairs
everywhere blackish at base; terminal half of fur on back and sides
cartridge buff; beneath, whitish. Feet, ears, wings, and tail nearest
avellaneous of Ridgway’s Color Standards; wing and tail membranes
faintly edged with whitish. A specimen from Carroll Creek, Inyo
County, is very similar in coloration, save that the ears, feet, and
wing and tail membranes are nearest clove brown. Two specimens
(coll. U. S. Nation. Mus.) from Colonia Lerdo, on the Colorado River,
below Yuma, in Sonora, Mexico, agree closely in coloration with the
Fort Mohave specimen; while a specimen (no. 138556, coll. U. S.
Nation. Mus.) from Rancho San Antonio, Lower California, at the
west base of the San Pedro Martir Mountains, varies in the direction
of specimens of Myotis ywmanensis sociabilis from Fort Tejon, Kern
County, California.
SkulltIntermediate in size between that of Myotis californicus
and Myotis lucifugus (total length, about 13.4 millimeters). Rostrum
and interorbital constriction broader in ywmanensis than in califor-
nicus. Skull of ywmanensis smaller and more slender than skull of
lucifugus, and brain-case more inflated in frontal region.
Measurements—The example of M. y. yumanensis (no. 16306,
Mus. Vert. Zool.) taken at Carroll Creek, Sierra Nevada Mountains,
Inyo County, September 9, 1911, measures in millimeters as follows:
total length, 84.0; tail vertebrae, 37.0; tibia, 14.8; foot, 9.5; forearm,
34.4; greatest length of cranium, 13.4; zygomatic breadth, 8.0; breadth
of brain-ease, 7.0; interorbital constriction, 3.8.
Synonymy and History.—tThis bat was described under the name
Vespertilio yumanensis by H. Allen (1864, pp. 58-59) from material
obtained at Fort Yuma, Imperial County, California. According to
Miller (1897), p. 67) this bat is also the Vespertilio macropus and
Vespertilio nitidus macropus of the same author, Dr. Harrison Allen.
Distribution —Muiller (1912, p. 56) gives the range of the species
as the Austral zones and lower edge of the Transition zone from the
southwestern United States to San Luis Potosi and Michoacan, Mexico.
The distribution in California is given by J. Grinnell (1913), pp.
276-277) as follows: Lower and Upper Sonoran zones throughout
southern California, both east and west of the desert divides; north
ro
1918] Grinnell: A Synopsis of the Bats of California 275
through Owens Valley and through the San Joaquin and Sacramento
valleys, at least as far as Oroville, Butte County. But this includes
also the range of the next described subspecies. A recent examina-
tion of the material in the Museum of Vertebrate Zoology showed the
existence of a previously unnamed race, Myotis ywmanensis sociabilis
MYOTIS YUMANENSIS YUMANENSIS
MYOTIS YUMANENSIS SOCIABILIS |
MYOTIS YUMANENSIS SATURATUS
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA
Fig. J. Map showing stations of occurrence in California of Myotis ywman-
ensis yumanensis, Myotis ywmanensis sociabilis, and Myotis ywmanensis saturatus,
as established by specimens examined by the author.
(H. W. Grinnell, 1914, pp. 318-319), west of the desert divides, thus
restricting the range of the typical subspecies, Myotis yumanensis
yumanensis, to the arid Colorado and Mohave deserts and north
through the Inyo region, probably altogether within the Lower
Sonoran zone. (See map, text-fig. J.)
276 University of California Publications in Zoology [Vou.17
Specimens Examined.—But a single example of the Yuma bat from
California has been examined by the writer: Carroll Creek, Inyo
County, 1.
Natural History—The writer has found nothing published upon
the habits of the Yuma bat.
Myotis yumanensis sociabilis H. W. Grinnell
Tejon Bat
Myotis yumanensis, Miller (1897b, pp. 66-68), part. Description; dis-
tribution.
Myotis yumanensis, Elliot (1901, p. 403), part. Diagnosis; distribution.
Myotis yumanensis, Elliot (1904a, pp. 318-319), part. Occurrence at Fort
Tejon.
Myotis yumanensis, Elliot (1904b, pp. 576-577), part. Diagnosis; dis-
tribution.
Myotis yumanensis, Elliot (1905, p. 475), part. Diagnosis; distribution.
Myotis yumanensis, Stephens (1906, p. 267), part. Diagnosis; distri-
bution.
Myotis yumanensis, Elliot (1907, pp. 501-502), part. Localities of cap-
ture in California.
Myotis lucifugus longicrus, J. Grinnell (1908, p. 158), part. San Bernar-
dino Mountains.
Myotis yumanensis yumanensis, Miller (1912, p. 56), part. General range.
Myotis yumanensis yumanensis, J. Grinnell (1913b, pp. 276-277), part.
Range in California.
Myotis yumanensis yumanensis, Grinnell and Swarth (1913, pp. 380-381).
Record of capture in the San Jacinto Mountains.
Myotis ywmanensis sociabilis H. W. Grinnell (1914, pp. 318-319). Orig-
inal description; type locality, Fort Tejon, Kern County, California.
Diagnosis—Similar in general characters to Myotis yumanensis
yunanensis (H. Allen) and M. y. saturatus Miller, but intermediate
in color between these two forms.
Description—F ur distributed as in M. y. yumanensis; on middle
of back averages about six millimeters in length. Hairs everywhere
clove brown at bases; distal half of fur on dorsal surface wood
brown; fur below light buff, with darker bases of hairs showing
through. On throat, sides and chin the color varies toward warm
buff; ears olive brown; feet, wings and tail-membranes clove brown.
The young are darker and grayer throughout, entirely lacking the
buffy tint of the adults. Specimens of M. y. sociabilis from the San
Bernardino Mountains show strong superficial resemblance to the
smaller individuals among a series of Myotis longicrus interior Miller
from the same locality. The longer tibia of the latter species, how-
ever, together with the slightly greater size of skull and the more
elevated occipital region, serves to allocate individuals.
Measurements——A series of five adult males of M. y. sociabilis
averages in millimeters as follows: Total length, 81.4 (extremes
1918] Grinnell: A Synopsis of the Bats of California 277
75.0-87.0) ; tail vertebrae, 33.3 (30.0-37.0) ; tibia, 15.2 (15.0-16.0) ;
foot, 8.1 (7.0-10.0) ; forearm, 34.6 (32.9-35.3); greatest length of
eranium, 13.7 (13.5-14.4) ; zygomatic breadth, 8.4 (8.0-8.6) ; breadth
of brain-case, 7.1 (6.7—7.6) ; interorbital constriction, 3.8 (3.5-3.9).
Ten adult females from Fort Tejon, Kern County, average in milli-
meters as follows: Total length, 81.9 (extremes, 76.0—-85.0) ; tail
vertebrae, 36.6 (31.0-87.0) ; tibia, 14.7 (13.5-16.0) ; foot, 8.9 (8.0—
10.0); forearm, 34.2 (83.5-35.0) ; greatest length of cranium, 13.8
(13.4-14.2) ; zygomatic breadth, 8.1 (7.8-8.3) ; breadth of brain-case,
7.2 (6.77.3) ; interorbital constriction, 3.7 (3.5-4.0).
MEASUREMENTS IN MILLIMETERS OF TEN MALES oF Myoris YUMANENSIS
SOCIABILIS H. W. GRINNELL, FROM CALIFORNIA
z 2 E
Mus. ge as 5 3 z
no. Age a a a ey Lem
69617 juv. 71.0 30.0 12.6 8.0 30.0
6639" juv. 82.0 35.0 16.0 9.0 34.5
6645' = juv. 80.0 36.0 15.0 8.0 33.7
6662" juv. 67.0 31.0 12.8 8.0 Chile: Ween eee Ae
6636 juv. 82.0 36.0 14.5 8.0 BBY 13.2 7.9 6.7 3.5
6669* ad. 79.0 32.0 15.5 35.2 14.0 oe
6665° ad. 83. 30.0 15.0 35.0 14.4 a2 7.4 3.8
66679 — ad. 87.0 37.0 16.0 7.0 35.3 13.9 8.6 7.6 3.9
6666" ~~ ad. 83.0 35.0 15.0 8.0 35.0 14.1 8.6 7.5 a
18485* sad. 75.0 32! Ole hese 10.0 32.9 13.5 8.0 7.0 3.8
1From Fort Tejon, Kern County.
?From South Fork Santa Ana River, San Bernardino Mountains.
’From Bluff Lake, 7500 ft., San Bernardine Mountains.
4From Chambers Ravine, 4 miles north of Oroville, Butte County.
MEASUREMENTS IN MILLIMETERS OF TEN ADULT FEMALES OF MYOTIS YUMANENSIS
SOCIABILIS H. W. GRINNELL, FROM Fort TEJoN, Kern County, CALIFORNIA
no. aT =) a 4 = wo NT
6660 85.0 36.0 14.0 9.0 34.0 13.8 8.3
6642 81.0 36.0 14.6 9.0 33.8 3.7 8.2
6629 76.0 31.0 15.0 9.0 35.0 13.4 7.8
6624 81.0 34.0 13.5 8.0 33.5 MAW Be
6628 82.0 35.0 15.0 8.0 34.5 14.0
5149 82.0 37.0 15.0 10.0 34.3 13.8 8.0
6618 83.0 36.0 14.2 9.0 34.8 14.0 8.0
5157 82.0 36.0 14.7 9.0 34.0 13.9
5158 84.0 36.0 16.0 Oa) BBY 14.2 8.3
6616 83.0 34.0 15.0 9.0 34.8 14.0
278 University of California Publications in Zoology [Vou.17
Distribution—The distribution of this bat cannot be stated with
confidence without much further field work. It appears to occupy
a geographic position intermediate between that of M. y. ywmanensis
and M. y. satwratus, namely the semi-arid Transition and Sonoran
zones in California west and north of the southeastern deserts. (See
map, text-fig. J.)
Specimens Exanined.—Total number 69, from the following local-
ities in California: Butte County: Chambers Ravine, four miles north
of Oroville, 1; Glenn County: Winslow, five miles west of Fruto, 1;
Kern County: Fort Tejon, 61; Buttonwillow, 1 (Calif. Acad. Sei.) ;
San Bernardino County: Bluff Lake, 7500 feet altitude, 3; Bear
Lake, 6700 feet altitude, 1; South Fork Santa Ana River, 8500 feet
altitude, 1.
Natural History.—July 21 to 25, 1904, J. Grinnell collected sixty-
one bats of this species in and about the half-ruined buildings of old
Fort Tejon, Kern County. Of the specimens secured, thirty-three
proved to be adult females, twelve young females and sixteen young
males. There were no adult males in the lot taken. Some of the
young males are equal in length to fully adult males from other
localities, but their immaturity is attested by the swollen and con-
spicuous finger joints as well as by the appearance of the pelage. It
is possible that with the approach of summer the full-grown males
leave the colony and forage singly at higher elevations.
An example of this species of bat (no. 6669, Mus. Vert. Zool.)
was taken at an elevation of 8500 feet on the South Fork of the
Santa Ana River in the San Bernardino Mountains. This was the
highest station for any species of bat secured in those mountains.
J. Grinnell (1908, p. 158) erroneously records the taking of this
specimen under the name Myotis lucifugus longicrus.
Myotis yumanensis saturatus Miller
Miller Bat
Myotis ywmanensis saturatus Miller (1897b, p. 68). Original description ;
type locality, Hamilton, Skagit County, Washington.
Myotis yumanensis saturatus, Stephens (1906, p. 267), part(?). Diagnosis;
distribution.
Myotis yumanensis saturatus, J. Grinnell (1913b, p. 277), part. Range
in California.
Dagnosis—Similar to Myotis ywmanensis yumanensis and M. y.
sociabilis, but fur longer, and color darker; general color of back
bister.
1918] Grinnell: A Synopsis of the Bats of California 279
Description.—A specimen (no. 11844, Mus. Vert. Zool.) taken by
F. Stephens at Cuddeback, Humboldt County, California, September
16, 1910, presents the following coloration: Back, nearest bister ; lower
surface, drab, darkest on chin, throat and sides; membranes, chaetura
black. Fur 6 to 7 millimeters long on middle of back.
Measurements.—The above specified example measures: Total
leneth, 84.0; tail vertebrae, 35.0; tibia, 14.1; foot, 8.0; forearm, 35.0;
ereatest length of cranium, 13.9.
Synonymy and History.—This species was described, under the
name Myotis ywmanensis saturatus, by Miller (1897), p. 68). The
type specimen was taken at Hamilton, Skagit County, Washington.
Distribution—Myotis ywmanensis saturatus is the dark northwest
coast form of the Yuma bat. Miller (1912, p. 56) gives the range
of this race as the Transition Zone in Oregon, Washington, and British
Columbia. Its distribution in California is given by J. Grinnell
(19136, p. 277) as the Transition and Boreal zones in extreme north-
western California, west to Cuddeback, Humboldt County, and east
to Mount Shasta. The specimens upon which this statement of the
Californian distribution of the Miller bat was based have been exam-
ined by the present writer who finds that the examples from Mount
Shasta are Myotis lucifugus altipetens, as are also the specimens from
the same locality recorded under the name saturatus by C. H. Merriam
(1899, p. 89). [See under Myotis lucifugus altipetens, p. 264.] This
leaves Eureka and Cuddeback, in Humboldt County, the only verified
record stations, within the state. This bat will probably be found to
occur at many points within the extreme northwestern portion of
California, where humid conditions prevail. (See map, text-fig. J.)
Specimens Examined.—The writer has examined but two speci-
mens of the Miller bat from within the state of California. They
were taken at Eureka (no. 11854, Mus. Vert. Zool., August 2, 1910)
and Cuddeback (see under Description, above), both localities being
in Humboldt County.
Natural History.—Nothing has been recorded concerning the
natural history of Myotis ywmanensis saturatus as occurring in Cali-
fornia.
Myotis californicus californicus (Audubon and Bachman)
Little California Bat
Vespertilio californicus Audubon and Bachman (1842, pp. 285-287). Orig-
inal deseription; type locality, ‘‘California.’’
Vespertilio nitidus H. Allen (1862, pp. 247-248). Original description;
no type designated, but specimens listed from Monterey, California, and
Fort Steilacoom, Washington.
280 University of California Publications in Zoology [Vou.17
Vespertilio nitidus, H. Allen (1864, pp. 60-62, fig. 57-59), part. Deserip-
tion; distribution.
V [espertilio]. nitidus, Cooper (in Cronise, 1868, p. 442). Listed as oceur-
ring in California.
V [espertilio]. obscurus, Cooper (in Cronise, 1868, p. 422)(?). Listed as
oceurring in California.
Vespertilio nitidus, Dobson (1878, pp. 318-319, pl. 29, fig. 7), part. De-
scription; general distribution (includes ‘‘California’’).
Vespertilio nitidus, H. Allen (1894, pp. 94-104, pl. 12), part. Desecrip-
tion; distribution.
Myotis californicus, Miller (1897b, pp. 69-72), part. Description; dis-
tribution.
Myotis californicus, Merriam (1899, p. 89). Occurrence on Mt. Shasta.
Myotis californicus caurinus, Stone (1904a, p. 579). Record of occurrence
on Mt. Sanhedrin.
Myotis californicus, Elliot (1901, pp. 403-404), part.
tribution.
Myotis californicus, Elliot (1904b, pp. 578-579). Diagnosis; distribution.
Myotis californicus, Stephens (1906, p. 266), part. Diagnosis; distribution ;
habits.
Myotis californicus, Miller (1912, p. 56), part. Range.
Myotis californicus, J. Grinnell (1913b, p. 277), part. Distribution in
California.
Diagnosis; dis-
Diagnosis —Total length, 74 to 83 millimeters ; forearm, 30 to°31.6;
foot weak, slender, and less than half leneth of tibia; calear about
as lone as free edge of interfemoral membrane, very slender, edge
ss with a distinct keel, lobed at tip. Free
\ border of uropatagium naked. Ears
AS) aC oe >)
L
Fig. K. Side view of head
of Myotis californicus califor-
nicus (drawn from specimen
no. 21875), X 1.00, showing
erect and acutely pointed tra-
gus.
Fig. L. Side view of head
of Pipistrellus hesperus hes-
perus (drawn from specimen
no. 10772), X 1.00, showing
reaching just beyond tip of nose.
Tragus pointed at tip and directed
upward, not forward (see text-fig. K).
Wing membrane attached at bases of
toes. Fur on back distinetly darker at
base than at tip; general color of back
mummy brown. (See pl. 18, fig. 17).
Description: Head.—Kar moderately
long for the genus (12 to 14 millimeters
in height from meatus), and when laid
forward reaching from 1 to 3 miulli-
meters beyond tip of nose. Anterior
border of auricle straight or slightly
curved tragus with blunt tip.
convex at base, then strongly convex to
a pot somewhat beyond middle, thence straight or even a little con-
cave to rounded tip; posterior border coneave from tip to a point
slightly below middle, thence convex to basal notch; basal lobe
strongly developed and notched on its lower border. Tragus straight
and tapering, slightly more than half height of ear (text-fig. K).
Limbs and Membrancs.—All membranes thin and delicate. Wings
attached at bases of toes. Feet small and weak, and usually less
than half length of tibia. Calear slender and usually terminating
in a distinct lobe; posterior border provided with a keel.
1918 | Grinnell: A Synopsis of the Bats of California 281
Pelage-——Fur everywhere full and soft, 5 to 7 millimeters in
leneth on middle of back. Basal third of dorsal surface of ear
furred; ventral (inner) side with fine, short hairs sparsely sprinkled
over entire surface. Wings naked save for a narrow strip of fur
along edge of body. Uropatagium furred on basal third, and on its
ventral surface fine scattered hairs nearly to tip of tail. Upper
surface of toes sparsely coated with short fine hairs.
Color—A specimen (no. 12981, Mus. Vert. Zool.) taken in the
Yosemite Valley, elevation 4000 feet, May 30, 1911, presents the fol-
lowing coloration: General color of back mummy brown. Hairs on
middle of back chaetura black on proximal 4 millimeters, then wood
brown for about 1 millimeter, and with a 2 millimeter tip of mummy
brown. On dorsal surface of interfemoral membrane and at bases
of ears the hairs lack the black bases. On ventral surface of body
the hairs are chaetura black at their bases (except those on the inter-
femoral membrane, which are entirely buffy brown), with the terminal
millimeter buffy brown (brightening toward mummy brown on sides
of body). Ears, wings, and interfemoral membrane blackish.
Miller (1897), p. 70) gives the typical color of this bat as ‘‘heht
yellowish gray, paler on the belly, the fur everywhere dark plumbeous
at base. Membranes, ears, lips, and muzzle blackish.’’ At this time
Miller recognized the existence of but one race of Myotis californicus
within the state of California, and his color description is evidently
taken from examples of the pale desert race, Myotis californicus
pallidus.
Skull—Small as compared with skulls of other California species
of Myotis (12.0 to 13.3 millimeters in greatest length), and delicately
formed. Brain-ease rounded and forehead sloping gently, forming a
comparatively deep saddle between brain-case and up-turned rostrum
(pl. 22, fig. 36). Interorbital constriction narrow (3.0 to 3.2 milli-
meters in width). Occipital elevation slight. Skull of same general
type as in Myotis evotis and M. thysanodes, but readily distinguished
by its much smaller size. Interorbital constriction (pl. 21, fig. 24)
much narrower in californicus than in yumanensis. In greatest
leneth of cranium californicus is only slightly smaller than orinomus,
but in lateral profile the ‘‘flattening’’ of the skull of oriznomus (pl. 22,
fig. 38) contrasts strongly with the saddle-shape in californicus (pl. 22,
fie. 836) as formed by the angular outline of the rostrum and forehead.
Measurements —Ten adult specimens of this bat average in milli-
meters as follows: total length, 77.6 (extremes, 74.0-83.0) ; tail verte-
brae, 34.6 (29.0-39.0) ; tibia, 13.2 (12.6-14.0); foot, 6.4 (5.5-8.0) ;
forearm, 30.8 (30.0-31.6) ; greatest length of cranium, 13.0 (12.4—
13.3) ; zygomatic breadth, 7.6 (7.1-8.1); breadth of brain-ease, 6.7
(6.4-7.0) ; interorbital constriction, 3.1 (3.1-3.2).
Synonymy and History—tThe little California bat was described
by Audubon and Bachman (1842, pp. 285-287) under the name
Vespertilio califormecus. The deseribers state: ‘‘We have obtained
but a single specimen, which was captured at California.’’ Miller
(18976, pp. 21-22) has shown the description given by Audubon and
Bachman to be applicable to Myotis californicus alone among the
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OOL6T
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GQ giody Nay dO SHULANITTIIP, NI SLNANAYOASVafL
1918 | Grinnell: A Synopsis of the Bats of California 283
various small bats known to oceur within the state of California. He
does not, however, indicate to which of the races of californicus the
name may be taken to apply. According to H. Allen (1894, p. 94),
the type is lost. I have here applied the name californicus to the
race occupying that portion of California north of about latitude
36 degrees and west of the desert divides. H. Allen (1862, pp.
247-248) describes five specimens of californicus under the new name
Vespertilio nitidus, concluding that the description given by Audubon
and Bachman is not sufficiently clear to identify the animal. Since
the specimen first mentioned by Allen came from Monterey, Califor-
nia, Miller (1897), p. 69) lists nitidus as a synonym of californicus.
Lyon and Osgood (1909, p. 272), however, consider, for reasons which
they state, that the type should be regarded as having come from
Fort Steilacoom, Washington (where four of the five specimens were
secured). In the latter case it would apply to the race of californicus
now known as Myotis californicus cawrinus Miller. I have accepted
Miller’s ruling, whereby ‘nitidus becomes a synonym of californicus.
I have examined the five specimens from Mount Sanhedrin which
were referred to the race caurinus by Rehn (in Stone, 1904a, p. 579)
and find them to belong rather to M. c. californicus, as here under-
stood.
Remarks.—In a series of fifty skins of the races califormecus and
quercinus there is a lack of the uniformity of coloration which so
clearly marks the race pallidus, and yet the distinctness of the two
first-mentioned races cannot be questioned when the mass coloration
is considered. A series of seventeen skins of Myotis californicus cali-
fornicus, including both adults and young, taken during July at the
one locality, Fyffe, Eldorado County, shows extremes, the paler of
which is indeed lighter colored than the darkest quercinus; but the
mean places the entire series with californicus, the latter as repre-
sented by a series from west-central California. Stephens (1906,
pp. 266-267), a field naturalist of long experience in California,
considers bats of the Myotis californicus group migratory and states
that ‘‘a few Bats winter in the Colorado Desert; these appear to be
intermediate between pallidus and californicus.’’ This statement by
Stephens calls to mind certain lines from the paper by Murphy and
Nichols (1918, p. 6) on Long Island bats:
... We may attribute to the Silver-haired Bat and other bats, a type of migra-
tion analogous to that of many birds, in which the individuals of a species within
a given breeding range move southward in fall, only to be replaced by winter
284 University of California Publications in Zoology [Vou.17
residents of the same species coming from a more northerly faunal area, On
such a hypothesis, a counter tendency in spring would cause a northward flight
of the species as a whole, until each group had reached its native habitat.
The two darkest skins in the Museum series referred to Myotis
californicus quercinus are autumn specimens, one (no. 6952) having
4 MYOTIS CALIFORNICUS CALIFORNICUS 7
@ MYOTIS CALIFORNICUS QUERCINUS
@ MYOTIS CALIFORNICUS PALLIDUS
ear:
ye
DISTRIBUTION MAP
MUSEUM OF YERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA,
al ma = = = |}
Fig. M. Map showing stations of occurrence in California of Myotis cali-
fornicus californicus, Myotis californicus quercinus, and Myotis californicus pal-
lidus, as established by specimens examined by the author,
been taken at Pasadena, September 24, and the other (no. 2803) in
the Cuyamaca Mountains of San Diego County, August 18. It is
possible that these examples are fall migrants of M. c. califormcus.
Much more field work remains to be done before the status and dis-
tribution of these subspecies can be stated with confidence.
1918] Grinnell: A Synopsis of the Bats of California 285
Distribution.—M yotis californicus californicus, as here restricted,
occupies that portion of California north of about latitude 36 degrees
and west of the Sierran divides. It appears to inhabit exclusively the
Upper Sonoran and Transition zones. (See map, text-fig. M.)
Specimens Examined.—The writer has examined 81 specimens
(mostly skins with skulls) of M. c. californicus from the following
localities in California: Mendocino County: Laytonville, 1; Sherwood,
10 (Calif. Acad. Sei., 9; Mus. Vert. Zool., 1) ; Willits, 1; Mt. Sanhed-
rin, 5 (Acad. Nat. Sci. Phila.) ; 3 miles west Mt. Sanhedrin, 1; Glenn
County : Winslow, 1; Sonoma County : near Cazadero, 3; near Guerne-
ville, 2; Colusa County: Snow Mt., 1 (Stanford Univ.) ; Solano
County: near Vacaville, 3; Contra Costa County: near Walnut
Creek, 4; Monterey County: Monterey, 5; Placer County: Blue
Canon, 1; Dutch Flat, 2; Eldorado County: Fyffe, 28; Limekiln, 1;
Michigan Bluff, 2; Placerville, 1 (Stanford Univ.) ; Mariposa County :
Yosemite Valley, 2; El Portal, 2; Pleasant Valley, 4; Coulterville, 1.
Natural History—When overtaken by a storm in the Yosemite
Valley on May 30, 1911, the writer sought refuge in a rocky cavern.
A tiny fire was built for comfort and in a few moments a little Cali-
fornia bat dropped from a erevice in the roof, evidently overcome
by the smoke. The specimen proved to be an adult male and is now
no. 12981 (Mus. Vert. Zool.).
A female (no. 18487, Mus. Vert. Zool.), with mammae functional,
was taken July 2, 1912, three miles west of Vacaville, Solano County.
Among a series of twenty-eight little California bats in the
Museum of Vertebrate Zoology, secured by Joseph Dixon at Fyffe,
Eldorado County, during the latter part of July, 1916, there are
adults and young of both sexes. The young are nearly full-grown
and vary in weight from 2.8 grams to 3.1 grams. The largest of the
adult males has a length of 80 millimeters and weighs 3.7 grams, while
among the females the greatest length is 80 millimeters, with a weight
of 5 grams.
Myotis californicus quercinus II. W. Grinnell
Oak Foliage Bat
Myotis californicus, Elliot (1904a, p. 319), part. Occurrence at Fort Tejon
and Mt. Whitney, California.
Myotis califernicus, Elliot (1907, pp. 502-503), part. Mt. Whitney and
Fort Tejon.
Myotis californicus, J. Grinnell (1908, p. 158). Occurrence in the San
Bernardino Mountains.
bo
(o3)
for)
University of California Publications in Zoology {|Vou.17
Myotis californicus californicus, Grinnell and Swarth (1913, p. 381).
Occurrence in the San Jacinto Mountains.
Myotis californicus quercinus H. W. Grinnell (1914, pp. 317-318). Orig-
inal description; type locality, Seven Oaks, San Bernardino County.
Diagnosis—Similar to Myotis californicus californicus (Audubon
and Bachman) and Myotis californicus pallidus Stephens, but inter-
mediate in color between these two forms. Prevailing tone of color
on back, cinnamon.
Description.—Kars, feet and fur as in M. ce. californicus. Mem-
branes and bases of hairs everywhere as in californicus. Terminal
portions of fur on back, glossy cmnamon, this color extending down
onto sides. Terminal portions of hairs on lower surface of body hght
buff, rather than buffy-brown as in californicus, or pale cartridge-
buff as in pallidus.
Measurements.—A series of ten examples of M. c. quercinus from
southern California averages in millimeters as follows: Total length,
81.6 (extremes, 77.0-83.0); tail vertebrae, 36.8 (31.0-41.0) ; tibia,
14.1 (12.5-15.0); foot, 6.0 (4.0-8.0); forearm, 31.9 (31.0-33.2) ;
greatest length of cranium, 13.1 (12.9-13.8) ; zygomatic breadth, 7.7
(7.4-8.0) ; breadth of brain-case, 6.8 (6.6—7.0) ; interorbital constric-
tion, 3.0 (2.9-3.3).
Synonymy and History—This form was always included under
the name M. c. califormicus, until described by the present writer
(1914; p. 317).
Distribution—tThe range of M. c. quercinus, as so far worked
out, occupies portions of the San Diegan faunal division of southern
California, and the Santa Barbara Islands. The life-zone is high
Upper Sonoran and low Transition. (See map, text-fig. M.)
Specimens Examined—The writer has examined 41 specimens
of Myotis californicus quercinus from the following loealities in Cali-
fornia: San Diego County: Cuyamaeca, 2; Julian, 5; Witch Creek, 2
(San Diego Soe. Nat. Hist.) ; Dulzura, 3 (Amer. Mus. Nat. Hist.) ;
Santa Cruz Island: Friar’s Harbor, 3; San Clemente Island, 4; Santa
Catalina Island, 1 (coll. F. W. Koch) ; San Bernardino County : Seven
Oaks, 2; Bear Lake, 1; South Fork Santa Ana River, 2; San Jacinto
Mountains, Riverside County: Kenworthy, 1; Schain’s Ranch, 1; Los
Angeles County: Los Angeles, 1 (Amer. Mus. Nat. Hist.) ; Pasadena,
1; Ventura County: Matilija, 1; Mount Pinos, 2; Tulare County:
Trout Creek, 2; Kern County: San Emigdio, 7.
Remarks.—The specimens listed from Santa Cruz, San Clemente,
and Santa Catalina islands, while slightly darker than typical quer-
cinus, are still nearer to this form than to M. c. californicus.
Natural History —tIn the San Bernardino Mountains, J. Grinnell
(1908, p. 158) found these bats in the Transition Zone, where they
were flitting close about the foliage of oaks and pines at late twilight.
287
f the Bats of California
: A Synopsis o
Grinnell
1918}
MEASUREMENTS IN MILLIMETERS OF TEN EXAMPLES OF
Mus.
no.
6671
6940
6673
6675
6939"
2780
1894
2781
16301
6672
1 Type.
* Probably a mistake; in the dried skin the foot measures 6 millimeters,
Sex
4010 Q,0Q,4040 +0 1+04040
Locality
Mt. Pinos, 6500 ft., Ventura Co.
Friar’s Harbor, Santa Cruz Isl.
Friar’s Harbor, Santa Cruz Isl.
South Fork Santa Ana, 6200 ft., San Ber-
nardino Mts.
Seven Oaks, 5000 ft., San Bernardino Mts.
Julian, San Diego Co.
Kenworthy, 4500 ft., San Jacinto Mts.
Julian, San Diego Co.
Trout Creek, 6000 ft., Tulare Co.
Bear Lake, 6700 ft., San Bernardino Mts.
Myorts CALIFORNICUS
Tibia
13.8
14.0
15.0
14.0
14.2
13.7
14.5
14.6
15.0
12.5
Foot
6.0
5.0
4.0
7.0
6.0
7.0
6.0
8.0
5.0
6.0
Forearm
length of
eranium
Greatest
Zvgomatie
Go breadth
a
Sw
em
Breadth of
brain-case
QUERCINUS H, W. GRINNELL, FROM CALIFORNIA.
constriction
co w pro Interorbital
Scot
3.2
3.0
288 University of California Publications in Zoology [Vou.17
The breeding season of M. c. quercinus is indicated by the follow-
ing data, taken from specimens in the Museum of Vertebrate Zoology.
No. 6675, female with one embryo taken July 3, 1906, on the South
Fork of the Santa Ana River, San Bernardino Mountains, 6200 feet
altitude; no. 6939, female with one embryo, taken July 8, 1905, at
Seven Oaks, San Bernardino Mountains, 5000 feet altitude.
Myotis californicus pallidus Stephens
| Little Pallid Bat
Myotis californicus, Miller (1897b, pp. 69-72), part. Specimens listed
from localities in southeastern California.
Myotis californicus pallidus Stephens (1900, p. 153). Original deseription ;
type locality, Vallecito, San Diego County, California.
Myotis californicus pallidus, Elliot (1901, p. 405). Diagnosis.
Myotis californicus pallidus, Miller and Rehn (1901, p. 256). Type
locality.
Myotis californicus pallidus, Elliot (1904a, p. 319). Record stations for
California: Mesquite Valley and Panamint Mountains, Inyo County.
Myotis californicus pallidus, Elliot (1904b, p. 579). Geographic distri-
bution; general characters.
Myotis californicus pallidus, Elliot (1905, p. 477). Geographic distri-
bution.
Myotis californicus pallidus, Stephens (1906, pp. 266-267). Description;
distribution.
Myotis californicus pallidus, Elliot (1907, p. 503). Repeated records. .
Myotis californicus pallidus, Lyon and Osgood (1909, p. 291). Nature and
location of type.
Myotis californicus pallidus, Miller (1912, p. 57). Nominal.
Myotis californicus pallidus, J. Grinnell (1913b, pp. 277-278). Range
in California.
Myotis californicus pallidus, J. Grinnell (1914, pp. 265-266). Occurrence
on the Colorado River; habits.
Diagnosis —Similar to Myotis californicus californicus and M. c.
quercinus, but averaging slightly smaller, with skull smaller (pl. 21,
fig. 25), and color lighter; general tone of back light buff.
Description: Color—A dry skin of Myotis californicus pallidus
(no. 7350, Mus. Vert. Zool.) taken by F’. Stephens at the type locality,
Vallecito, San Diego County, presents the following coloration: Basal
3 millimeters of fur on back, chaetura black, followed by a band of
cartridge buff 1 millimeter wide; terminal 3 millimeters, light buff.
On dorsal surface of interfemoral membrane the hairs lack the black
bases. Fur on ventral surface pale cartridge buff, with blackish bases
of hairs showing plainly. Ears, wings and interfemoral membrane
vary in color from drab to hair brown.
Measurements.—A series of five males of M. c. pallidus averages
in millimeters as follows: total length, 81.4 (extremes, 77.0-85.0) ;
tail vertebrae, 40.6 (38.0-44.0); tibia, 13.4 (13.0-14.1); foot, 7.2
1918 | Grinnell: A Synopsis of the Bats of California 289
(6.0-8.0) ; forearm, 30.5 (29.5-31.5); greatest length of cranium,
12.8 (12.8-12.9) ; zygomatic breadth, 7.5 (7.3-7.9) ; breadth of brain-
case, 6.6 (6.46.8) ; interorbital constriction, 2.9 (2.8-3.1).
MEASUREMENTS IN MILLIMETERS OF TEN ADULTS OF MYOTIS CALIFORNICUS
PALLIDUS STEPHENS, FROM CALIFORNIA
ah = S ~ 5 SS ate)
Mus. . 35 Be 2 2 A se PH =9
no. Sex = (y= a om om N77 [=a] aha
10700! fof 81.0 40.0 13.0 6.0 ales) esta 7.9 6.8 3.0
166567 ret 82.0 42.0 13.0 8.0 30.0 12.8 ed) 6.7 3.1
17787° rot 77.0 38.0 13.0 7.0 31.5 12.8 7 hes) 6.4 2.9
19278* Jb 85.0 44.0 14.1 8.0 30.0 12.9 wis 6.5 3.0
19280" ref 82.0 39.0 14.0 7.0 AOI) ae ae =r 2.8
10701" 2 75.0 35.0 14.0 60s, y) Sx 13.2 ae 6.3 2.9
16657" 2 81.0 39.0 14.5 7.0 31.5 13.0 aay 6.4 3.0
18724* Q 82. 41.0 13.6 7.0 BOLT 12.9 cece 6.4 2.9
7350" ic) 84.0 40.0 13.9 7.0 31.0 12.5 6.4 3.0
19279* 2 78.0 38.0 14.0 7.0 30.4 12.5 6.3 2.9
1From Colorado Kiver, opposite The Needles.
>From Vallecito, San Diego County.
* From Lone Pine Creek, 4500 ft., Inyo County.
‘From La Puerta Valley, San Diego County.
Synonymy and History.—Myotis californicus pallidus is the pale
desert form of californicus. It was described by Stephens (1900,
p. 153), from material secured at Vallecito, San Diego County.
Distribution—Lower Sonoran Zone on the Colorado and Mohave
deserts, and north in Owens Valley at least to Lone Pine. (See map,
text-fig. M.)
Specimens Examined.—Total number, 15, from the following
localities in California: Inyo County: Lone Pine Creek, 1; Mesquite
Valley, 1 (Field Col. Mus.); Panamint Mountains, 1 (Field Col.
Mus.) ; Kern County: Redrock Canon, 1; San Bernardino County :
Colorado River, 2; Imperial County: Pilot Knob, 1 (San Diego Soe.
Nat. Hist.) ; San Diego County: La Puerta Valley, 4; Vallecito, 3;
Borega Spring, 1 (U.S. Biol. Surv.).
Natural History.—In the spring of 1910 a party of collectors from
the Museum of Vertebrate Zoology took four specimens of this bat
on the Colorado River and were confident that they saw many others.
J. Grinnell (1914, pp. 265-266) states that the specimens secured
were all taken at late dusk, considerably later than the bulk appear-
ance of Pipistrellus hesperus. Instead of flying high against the sky,
as is the habit of the latter species, M. c. pallidus was almost always
290 University of California Publications in Zoology (Vou. 17
seen low over the bushes of the second bottom, or along shallow washes
between clumps of mesquite, seldom appearing above the skyline.
Stephens (1906, p. 266) states that a female little pallid bat cap-
tured on April 29 contained one small foetus.
Myotis orinomus Elliot
La Grulla Brown Bat
Myotis orinomus Elliot (1903, pp. 228-229). Original description; type
locality, La Grulla, San Pedro Martir Mountains, Lower California.
Myotis lucifugus longicrus, J. Grinnell (1908, p. 158), part. San Bernar-
dino Mountains.
Myotis orinomus, Grinnell and Swarth (1912, pp. 137-142). Occurrence
in California.
Myotis orinomus, J. Grinnell (1918), p. 278). Range.
Diagnosis —Similar in general characters to Myotis californicus
californicus, but size slightly larger (total length 79 to 88 milli-
meters), thumb much longer, color paler (tawny olive), cranium
flatter and rostrum much broader.
Description: Head—In general appearance similar to that of
Myotis californicus californicus but slightly larger. In a series of
ten dried skins the ear averages in height 11.5 millimeters from notch,
and the tragus 7.0 millimeters in height. Tragus tall, slender and
tapering, with tip bluntly rounded.
Limbs and Membranes——Wing membrane arises from near bases
of toes. Calear longer than free border of interfemoral membrane,
and distinctly keeled; in four of ten specimens examined it is lobed
at tip.
Pelage—Fur everywhere full and soft, and of a silky texture on
the back. Most hairs on back average about 6 millimeters in length,
but scattered among them is a more scanty growth of hairs about 8
millimeters in length. Wing furred only at extreme base, both above
and below. Uropatagium scantily haired on basal fifth, both above
and below; on its ventral surface short hairs are still more sparingly
scattered over the remaining area. Ear haired on basal fifth of
posterior surface ; also a very scanty growth of fine short hairs extends
over entire anterior surface. A few long hairs project from gland-
ular portions of face, and a row of downward projecting hairs extends
along upper lip.
Color.—While a small amount of individual variation in color is
apparent in the specimens of orinomus at hand, the series as a whole
bears a very close resemblance in color to a series of Myotis occultus
from the Colorado River. The hairs are everywhere chaetura black
at their bases. On the dorsal surface the color of the distal portion
of the fur varies from light sayal brown to tawny-olive. On the sides
the darker shade of the back fades gradually into the cartridge buff
of the lower surface. The membranes are a dark clove brown.
Skull—kRKasily distinguished from that of any other small Myotis
by reason of its decidedly flattened rostrum and brain-ease (see pl. 21,
fig. 26, and pl. 22, fig. 38). This flattening has tended to obliterate
291
ul
f the Bats of Californ
: A Synopsis o
Grinnell
1918]
Mus.
no,
2044
2045
2046
2047
2048
6941
16300
16304
16305
16303
Sex
+0 +0 +0 0 00,0, 0, 0,0,
MEASUREMENTS IN MILLIMETERS
Locality
Garnet Queen Mine, Santa Ros:
Garnet Queen Mine, Santa Rosa
Garnet Queen Mine, Santa Rosa
Garnet Queen Mine, Santa Rosa
Hemet Lake, San Jacinto Mts.
Doble, San Bernardino Mts.
Walker Pass, Kern Co.
Fay Creek, near Weldon, Kern
Fay Creek, near Weldon, Kern
Carroll Creek, Inyo Co.
oF TEN
Mts.
Mts.
Mts.
Mts.
Co.
EXAMPLES OF MyoTIS oRINOMUS ELLIOT, FROM CALIFORNIA
vertebrae
wr
oo 7
c)
Tibia
13.0
12.3
12.4
13.0
12.8
14.0
13.6
13.7
14.0
13.5
Foot
7.0
6.0
7.0
8.0
8.0
6.0
6.5
7.0
7.0
6.0
Forearm
34.6
32.5
34.1
33.0
34.3
Greatest
length of
cranium
oo o Zygomatie
= breadth
NO
co Breadth of
do ~=6brain-case
eo Interorbital
constriction
ray
292 University of California Publications in Zoology [Vou.17
the angle or ‘‘saddle’’ between the rostrum and brain-case, which
is shown so clearly in many forms of Myotis. The tooth-row (length
5.2 to 5.5 millimeters) is much longer in orinomus than in californicus.
Measurements.—Average and’ extreme measurements in milli-
meters of a series of ten M. orinomus are as follows: Five males:
total length, 82.38 (extremes 80.0-86.0); tail vertebrae, 38.4 (37.0—
40.0); tibia, 12.7 (12.8-13.0); foot, 7.2 (6.0-8.0); forearm, 32.1
(30.8-33.2) ; greatest length of skull, 13.6 (13.4-14.0). Five females:
total length, 86.2 (82.0-88.0) ; tail vertebrae, 41.0 (38.0-45.0) ; tibia,
13.8 (13.5-14.0); foot, 6.5 (6.0-7.0); forearm, 33.7 (32.5-34.6) ;
greatest length of cranium, 14.3 (13.9-14.7). A comparison of the
above figures will show that the females average somewhat larger
than the males.
Synonymy and History.—The La Grulla brown bat was described
by Elhot (1903, pp. 228-229) from three specimens obtained in the
San Pedro Martir Mountains, Lower California, two being taken at
La Grulla and one at Santa Eulalia. The occurrence of this bat in
California is first mentioned by Grinnell and Swarth (1912, pp.
138-141).
Distribution—Myotis orinomus has been recorded only from
northern Lower California and from southern California, where its
range may be defined as the high Upper Sonoran zone, in its semi-
arid portion, from the east slope of the Sierra Nevada near Owens
Lake south through the southern Sierras and coast ranges to the
Mexican line. (See map,-text-fig. O.)
Specimens Examined.—The writer has examined 17 specimens of
Myotis orinomus from the following localities in California: San
Diego County: Dulzura, 3 (Amer. Mus. Nat. Hist., 2; Acad. Nat. Sci.
Phila., 1); Jacumba, 1 (Amer. Mus. Nat. Hist.) ; Santa Ysabel; 1
(San Diego Soe. Nat. Hist.) ; Orange County: Trabuco Canon, Santa
Ana Mountains, 1; Riverside County: Garnet Queen Mine, Santa
Rosa Mountains, 4; Hemet Lake, San Jacinto Mountains, 1; San Ber-
nardino County: Doble, 1; Kern County: west slope Walker Pass, 1;
San Emigdio, 1; Fay Creek, 2; Inyo County: Carroll Creek, on east
slope Sierra Nevada, 1.
Natural History.—Nothing has been recorded of the habits of this
species.
Myotis evotis (H. Allen)
Little Big-eared Bat
Vespertilio evotis H. Allen (1864, pp. 48-50, figs. 42-44). Original
description; type locality fixed as Monterey, California (see Miller,
1897b, pp. 77-78).
Vespertilio evotis, Cooper (1868, p. 5). Distribution.
1918] Grinnell: A Synopsis of the Bats of California 293
V [espertilio]. evotis, Cooper (in Cronise, 1868, p. 442). Occurrence in
California.
Vespertilio evotis, Dobson (1878, p. 324). Description; general distri-
bution; one skin listed from ‘‘ California,’’
Vespertilio evotis, Bryant (1891a, p. 358). Nominal.
Vespertilio albescens evotis, H. Allen (1894, pp. 89-91), part. Descerip-
tion; distribution; localities of capture in California.
Myotis evotis, Miller (1897b, pp. 77-80). Description; distribution; speci-
mens listed from Inyo Mountains, Owens Lake, San Joaquin River, and
Twin Oaks.
Myotis evotis, Merriam (1899, p. 88). Occurrence on Mt. Shasta.
Myotis evotis, Elliot (1901, p. 406). Diagnosis; distribution.
Myotis evotis, Stone (1904a, p. 579). Record from Mt. Sanhedrin.
Myotis evotis, Elliot (1904a, p. 320). Occurrence on Mt. Whitney.
Myotis evotis, Elliot (1904b, p. 574). Diagnosis; distribution.
Myotis evotis, Stone (1904b, p. 587). Occurrence at Belmont.
Myotis evotis, Elliot (1905, p. 474). General distribution.
Myotis evotis, Stephens (1906, pp. 267-268). Diagnosis; distribution ;
migration.
Myotis evotis, Elliot (1907, p. 501). Record of occurrence in vicinity of
Mt. Whitney, California.
Myotis evotis, Miller (1912, p. 59). Range.
Myotis evotis, J. Grinnell (1913b, p. 278). Range in California.
Diagnosis.—Size large for a Myotis (length 75 to 93 millimeters) ;
forearm 35.7 to 38.4 millimeters; calear longer than free border of
interfemoral membrane; ear very long, when laid forward reaching
from 7 to 10 millimeters beyond snout.
Description: Head.—Kars large (19 to 20 millimeters in height
from meatus), thus easily distinguishing this species of Myotis from
all others occurring in California. Greatest width of ear shghtly
more than half its length. Anterior border of auricle convex from
base to a point slightly beyond the middle, thence straight, or nearly
so, to tip; posterior border slightly concave immediately below tip,
then gradually convex to base. Tragus long (10 to 13 millimeters),
slender, and pointed.
Limbs and Membranes.—Membranes thin and leht. Wings
attached at bases of toes; third and fifth metacarpals about equal in
length. Foot slightly less than half length of tibia. Calear slightly
lobed at tip, and longer than free border of uropatagium. Tail
slightly longer than forearm.
Pelage——¥ ur everywhere full and soft. Sides of face scantily
haired. Anterior border of ear scantily haired on basal fifth. Toes
slightly haired both above and beneath. Interfemoral membrane
naked save for basal fifth and a few scattering hairs along ‘‘nerves’’
and on free border.
Color.—Two skins from near Pasadena, Los Angeles County (nos.
6685, 6953), have ears and membranes dark clove brown; prevailing
tone of back pale honey yellow, with bases of hairs blackish slate; on
lower surface of body, outer portion of fur pale olive-buff. One
specimen (no. 12037) taken July 29, 1910, at Independence Lake,
294 University of California Publications in Zoology [Vou.17
Nevada County, is somewhat different in coloration, being paler and
erayer throughout, with ears and membranes blackish mouse gray.
Skull.—About 16 millimeters in length; slender and delicately
built (pl. 21, fig. 34, and pl. 22, fig. 46). Posterior margin of brain-
case rounded and occipital ridges but faintly defined. The only other
Californian species of this genus which have skulls similar in length
to that of evotis are Myotis velifer and M. thysanodes. Upon com-
parison it will be noted that in velifer the rostrum is heavily built,
the sagittal crest is distinct, and the posterior margin of the brain-
case is truncate, whereas in evotis the rostrum is lightly built, the
ridges are indistinct and the posterior margin of the brain-case is
rounded. The differences between the skulls of evotis and thysanodes
are not so tangible, and it is somewhat difficult to properly allocate
skulls of young individuals of these two species. But in adults
the brain-case of thysanodes is much more inflated than that of evotis,
the breadth of the brain-case in thysanodes is relatively greater, and
the rostrum is heavier.
Mcasurements——Measurements in millimeters of an adult female
(with one embryo) taken near Pasadena, May 29, 1904, are as fol-
lows: total length, 93.0; tail vertebrae, 43.0; tibia, 16.6; foot, 7.0;
forearm, 37.5; greatest length of cranium, 16.2; zygomatic breadth,
9.1; breadth of brain-case, 7.5; interorbital constriction, 3.7.
Synonymy and History.—Myotis evotis was described by H. Allen
(1864, pp. 48-50) under the name Vespertilio evotis. No type
locality was designated by that author, but Miller (1897), p. 78)
fixes Monterey, California, as the type locality, selecting one of the
localities given by H. Allen. In his monograph of 1894 (pp. 89-91),
Allen regards evotis as a race of Vespertilio albescens, and lists under
the same name a specimen of Myotis thysanodes (see Miller, 1897),
p. 80).
Distribution.—Given by Miller (1912, p. 59) as the Austral and
Transition zones from the Pacific coast to the eastern edge of the
Rocky Mountains; south to Vera Cruz, Mexico. In California the
range of Myotis evotis is not well defined as far as shown by the facts
yet available. It seems to comprise the Upper Sonoran and Transi-
tion zones from the Mexican line northwards as far as Mount Shasta;
west to the eastern boundary of Mendocino County, and east to Inde-
pendence Lake, Nevada County, and the Inyo Mountains, in Inyo
County. (See map, text-fig. N.)
Specimens Examined.—Total number, 22, from the following
localities in California: San Diego County: Dulzura, 2 (Amer. Mus.
Nat. Hist., 1; Acad. Nat. Sei. Phila., 1) ; Witch Creek, 2 (San Diego
Soe. Nat. Hist.) ; Twin Oaks, 1 (U.S. Biol. Surv.) ; San Bernardino
County: San Bernardino, 1 (San Diego Soe. Nat. Hist.) ; Los Angeles
County: near Pasadena, 2; North Fork San Gabriel River, 1 (Ll. A.
5
929!
ral
uu
f the Bats of Californ
ws O
Synopsi
A
Grinnell
1918 |
MEASUREMENTS IN MILLIMETERS OF SEVEN EXAMPLES OF MyorTiS EvoTIS (H. ALLEN), FROM CALIFORNIA
a a 3) "so
r= 5 z %28 es e:
ae) ae 3 = $ sue 8683 35
et Sex Locality ee S > Si é 2 ie = 3 et
6685 J Big Santa Anita Canon, near Pasadena 88.0? 36.0? 16.6 8.0 35.7 16.0 FAs 7.9
6953 2 Arroyo Seco, near Pasadena 93.0 43.0 16.6 7.0 37.5 16.2 9.1 7.5
12037 Jb Independence Lake, Nevada Co, 22 esses eee 16.8 ses 38.4 16.1 des 8.0
5759 Jb Wilts UMMUESS Joyakoeyeloy (Cf, ee 16.5 we DB Ote) ee esse He Sek
3283 3 Mt. Shasta, Siskiyou Co. 91.0 41.0 17.5 “as 3610) eee = eee
3284 Jb Mt. Shasta, Siskiyou Co. 90.0 43.0 16.1 ee =
160" 2 Pine Flat, N. Fork San Gabriel River,
Los Angeles Co. 75.0 GH) eres 9: Ob ees cot =
1Los Angeles County Museum of History, Science and Art.
> Interorbital
constriction
R=)
296 University of California Publications in Zoology |Vou.17
Co. Mus. Hist. Sci. and Art); Inyo County: Inyo Mts., 1 (U. 8.
Biol. Surv.) ; Owens Lake, 1 (U. 8S. Biol. Surv.) ; Madera County:
North Fork San Joaquin River, 1 (U.S. Biol. Surv.) ; Santa Clara
County: Palo Alto, 1 (Stanford Univ.) ; Saratoga, 1; San Mateo
bse “TRG as
Ce Te
@ MYOTIS EVOTIS
@ MYOTIS THYSANODES
4 MYOTIS VELIFER
Za, \Nih) 5 / m
oe ne
7 /\ve eee,
24 FS
COG A Nh ATS 3
~~
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNLA,
Fig. N. Map showing stations of occurrence in California of Myotis evotis,
Myotis velifer, and Myotis thysanodes, as established by specimens examined
by the author.
County: Pescadero Creek, 1; Belmont, 1 (Acad. Nat. Sci. Phila.) ;
Alameda County: Berkeley, 1 (U. C. Dept. Zool.) ; Eldorado County :
Mt. Tallac, 2 (Mus. Comp. Zool., 1; Mus. Vert. Zool., 1); Nevada
County: Independence Lake, 1; Mendocino County: Mt. Sanhedrin,
1 (Aead. Nat. Sci. Phila.) ; Siskiyou County: Mt. Shasta, 2.
1918] Grinnell: A Synopsis of the Bats of California 297
Natural History—lIt is probable that in California this species
is migratory and breeds chiefly in the Transition Zone. The only
specimens in the Museum of Vertebrate Zoology which are recorded
as secured in midsummer were taken in the Transition Zone. Stephens
reports securing specimens in San Diego County in spring and
autumn only.
As noted above, a female taken near Pasadena May 29 contained
a single embryo.
Myotis thysanodes Miller
Fringed Bat
Vespertilio albescens velifer (variety), H. Allen (1894, pp. 92-93). Dul-
zura (see Miller, 1897b, p. 80).
Vespertilio albescens evotis, H. Allen (1894, p. 90), part. Old Fort Tejon
(see Miller, 1897b, p. 80).
Myotis thysanodes Miller (1897b, pp. 80-85). Original description; type
locality, Fort Tejon.
Myotis evotis thysanodes, Elliot (1901, p. 406). Diagnosis; distribution.
Myotis thysanodes, Miller and Rehn (1901, p. 258). Type locality.
Myotis thysanodes, Elliot (1904b, pp. 572-573). Diagnosis; distribution.
Myotis thysanodes, Elliot (1905, p. 479). Distribution.
Myotis thysanodes, Stephens (1906, p. 268). Description; distribution.
Myotis thysanodes, Lyon and Osgood (1909, p. 273). Nature and location
of type.
Myotis thysanodes, Miller (1912,'p. 59). General range.
Myotis thysanodes, J. Grinnell (1913b, pp. 278-279). Range in California.
Diagnosis —Total length, 85 to 95 millimeters. Free border of
uropatagium thickened and densely haired.
Description —Largest Myotis found in California excepting Myotis
velifer. Easily distinguished from velifer by the much longer ears
which, when laid forward, reach from 3 to 5 millimeters beyond end
of nose. In an alcoholic specimen the ear measures 15 millimeters
from notch and the tragus 10 millimeters. Ear convex in outline
along anterior edge, bluntly rounded at tip and concave along upper
half of posterior margin, thence convex to the well-defined basal notch.
Tragus slender and straight, or nearly straight, along anterior edge
almost to tip, but convex just below bluntly rounded tip; posterior
border slightly concave just below tip, then convex; margin notched
just above the basal lobe.
Limbs and Membranes—-Wing membranes similar to those of
closely related species of Myotis, but interfemoral membrane heavier
and more leathery, and distinctly thickened on its free edge. Wings
arise just proximal to bases of toes. Calear distinct and slightly
lobed at tip. Foot slightly less than half length of tibia. Forearm
slightly longer than tail.
Pelage—Except for thickly haired border of uropatagium dis-
tribution of fur is similar to that of related species.
298 University of California Publications in Zoology {(Vou.17
Color.—As all the adult examples at hand are alcoholic the follow-
ing color description is taken from Miller (1897b, p. 81) :
The fur is everywhere light, dull, yellowish brown, distinctly paler ventrally,
the hairs everywhere dusky slate at base. The color is subject to considerable
individual variation in shade. The palest specimens are yellowish wood brown
inclining to clay color; the darkest specimens dull raw umber. The belly varies
from clear gray scarcely tinged with yellow to a strong yellowish gray, and in
other specimens to dull brownish gray. The exact shades are very variable and
impossible to describe accurately.
Skull—About 16 millimeters long, thus about equal to that of
Myotis velifer in total length, but quite distinct in general appearance
(pl. 21, fig. 35, and pl. 22, fig. 47). Rostral portion of skull relatively
slender (the diameter of rostrum, taken just posterior to the canines,
averaging a millimeter less in thysanodes than in velifer). Sagittal
crest well-defined in velifer, but indistinct in thysanodes. As pointed
out by Miller (1897b, p. 82) the posterior margin of the brain-case,
when viewed from above, is rounded in thysanodes, squarish, or
truneate, in velifer.
Measurements.—Miller (1897), p. 83) gives the average measure-
ments of a series of ten specimens of Myotis thysanodes from Fort
Tejon as follows: Total length, 87.0 millimeters; tail vertebrae, 37.0;
tibia, 17.6; foot, 8.0; forearm, 41.2; thumb, 6.3; longest finger, 69.2;
ear from meatus, 17.6; width of ear, 11.8; tragus, 10.5.
Synonymy and History.—This bat was described by Miller (1897),
pp. 80-85) under the name Myotis thysanodes. The type specimen
was obtained at Fort Tejon. Miller states that specimens of the
fringed bat were variously labeled or listed in his monographs by
H. Allen as ‘‘V[espertilio]. albescens velifer,’’ *‘V. subulatus,’’ “‘V.
albescens?,’’ and ‘‘V. albescens evotis.’’
Distribution —Given by Miller (1912, p. 59) as the Lower Sonoran
zone from near the southern border of the United States south to San
Luis Potosi and Michoacan, Mexico. The only Californian localities
of capture known to the writer besides those given by Miller (Fort
Tejon, Kern County, and Dulzura, San Diego County), are Limekiln
and Fyffe, Eldorado County. These four localities are all but one
well within the Upper Sonoran life-zone; Fyffe is in Transition. (See
map, text-fig. N.)
Specimens Examined.—Total number, 7, from the following locali-
ties in California: Kern County: Fort Tejon, 5 aleoholies (U. 8. Biol.
Surv.) ; Eldorado County: Limekiln, 1 (no. 24206, Mus. Vert. Zool.,
August 2, 1916) ; Fyffe, 1 (no. 24186, Mus. Vert. Zool., July 21, 1916).
Natural History—Miller (1897b, pp. 84-85) gives the following
account, furnished by Dr. T. S. Palmer, of the colony of bats from
which the type specimen of Myotis thysanodes was taken:
1918] Grinnell: A Synopsis of the Bats of California 299
In July, 1891, while one of the parties of the Death Valley Expedition was
collecting at Old Fort Tejon, California, several species of bats were observed.
The most abundant was a small Vespertilio [= Myotis|, which could be seen at
dusk flying about the oak trees near the old barracks in great numbers, and
passing in and out of the ruined buildings. A long two-story adobe building,
with the roof still intact, seemed to be the center of attraction, and about sundown
bats could be seen streaming forth from a window in one of the gables. On
the morning of July 5 an examination was made of the attic of this building,
and the bats were found clinging to the ridgepole and the rafters, literally by
thousands. Individuals of all ages, from recently born young to adults, were
hanging together in bunches as big as a bushel basket. Others found concealment
in cracks and crevices, but very few were flying about. Evidently the colony
had ocecupied.the attic for several years, but it was too dark to see whether more
than one species was present.
A sack was carried along under the ridgepole and specimens swept into it from
several of the larger bunches. In this way more than a hundred bats were
collected in a few minutes. As soon as they were disturbed they uttered a peculiar
squeaking note and flew about in a confused manner in their efforts to escape.
The sack was carried out under one of the oak trees and the specimens examined ;
160 had been captured, and of these 25 were preserved! and the remainder allowed
to escape. Some of the bats which had been given their liberty attempted to
fly back to their retreat, but dazed by the sunlight took refuge in the branches
of the nearest tree; others made no attempt to escape, except to crawl up the
trunks of the trees, where they remained until dark. Some of the young ones
failed to find their way back to the building, and remained about the spot for
several days.
In the summer of 1904 (July 19 to 26) J. Grinnell visited Fort
Tejon and collected bats in the same locality visited by Dr. Palmer
in 1891. Of the sixty-one examples of Myotis obtained on the later
date not one proved to be Myotis thysanodes.
Apparently bats of this species are not of wide distribution in
California, for among nearly a thousand specimens of bats collected
by the Museum of Vertebrate Zoology, only two have proven to be
Myotis thysanodes.
Genus Lasionycteris Peters
This genus includes the type species only, which is distributed
clear across northern North America and south nearly through the
United States.
Characters —Dental formula: i 72, e1!, pm22, m 22 = 36
aracters.—Denta ; — ea 3-3 3-30
SO Be atiqig © a) bilainy oe
Upper incisors conical; inner upper incisor slightly the longer,
and bicuspidate. Lower incisors subequal and crowded closely be-
tween canines; crowns of inner lower incisors four-lobed, those of the
outer lower incisors three-lobed. Canines both above and _ below
1 Sixteen proved to be Myotis thysanodes; the others were M. yumanensis.
300 University of California Publications in Zoology |Vou.17
simple, each with a distinct but rather small cmgulum. First upper
premolar minute; second slightly more than half height of canine.
First two lower premolars minute, being less than half the height
of the third.
Skull (pl. 21, fig. 33, and pl. 22, fig. 45) broad and somewhat
flattened; rostrum three-fourths width of brain-case, and strongly
coneave on each side back of nasal aperture.
Ear short, nearly as broad as long, and with a well-developed basal
lobe. Tragus short, straight, and bluntly rounded.
Lasionycteris noctivagans (Le Conte)
Silvery-haired Bat
V [espertilio|. noctivagans Le Conte (1831, p. 481). Original description;
no type locality designated.
Scotophilus noctivagans, H. Allen (1864, pp. 39-42, four figs. in text).
Description; listed from Fort Reading (in Shasta County).
Scotophilus noctivagans, Cooper (1868, p. 5). Distribution.
S[cotophilus|. noctivagans, Cooper (in Cronise, 1868, p. 442). Occurrence
in California.
Vesperugo noctivagans, Dobson (1878, pp. 238-239). Description; general
distribution (includes California).
Vesperugo noctivagans, C. H. Townsend (1887, p. 182). Record of speci-
men secured at Fort Reading by Dr. J. F. Hammond.
Vesperugo noctivagans, Bryant (1891la, p. 358). Nominal.
Lasionycteris noctivagans, H. Allen (1894, pp. 105-111, pls. 13-14). De-
scription.
Lasionycteris noctivagans, Miller (1897b, pp. 86-87). Description; dis-
tribution; listed from Nevada City and Nicasio.
Lasionycteris noctivagans, Elliot (1901, pp. 407-408, fig. 85). Diagnosis;
distribution.
Lasionycteris noctivagans, Stone (1904a, p. 579). Summer record from
Mt. Sanhedrin.
Lasionycteris noctivagans, Stephens (1906, pp. 268-269). Description;
distribution.
Lasionycieris noctivagans, Seton (1909, p. 1168). Map showing record
stations and hypothetical range.
Lasionycteris noctivagans, Miller (1912, p. 60). General range.
Lasionycteris noctivagans, J. Grinnell (1913b, p. 279). Range in Cali-
fornia.
Diagnosis ——Total length 92 to 107 millimeters; ears short and
rounded; color blackish chocolate, both above and below, many of the
hairs tipped with silvery white.
Description: Head.—Rostrum broad, two-thirds width of brain-
case; width between nostrils greater than distance from mouth to
top of rostrum; prominent glandular masses on each side of rostrum.
Ear short (height from meatus, 15 to 16 millimeters) and broad
(width, 11 to 14 millimeters) ; when laid forward reaching barely to
nostrils. Tragus a little less than half height of ear.
1918] Grinnell: A Synopsis of the Bats of California 301
Limbs and Membranes.—Wines moderately slender; third meta-
carpal slightly longer than fifth. Wing membranes attached to bases
of toes. Interfemoral membrane reaching to tip of tail. Foot slen-
der, somewhat compressed, and about half length of tibia. Calear
distinct and extending along edge of interfemoral membrane for about
a quarter of the distance from its origin to tip of tail.
Pelage.—Face nearly naked save for about a dozen slende hairs,
5 millimeters in length, which arise from the glandular masses on
sides of rostrum. Hair on top of head short, being only abont 2
millimeters in length; elsewhere on the body it is from 5 to 8 milli-
meters in length and very silky in texture. Kars naked, as also
the wings, save for a narrow strip of hair continuous with body fur.
Interfemoral membrane seantily haired over proximal half of its
dorsal surface; upon its ventral surface the hairs are even less plenti-
ful. Dorsal surface of toes well furred.
Color.—The fur varies from deep blackish chocolate, tipped with
silvery white, to a decided brownish, tipped with yellowish gray.
Seton (1909, p. 1167) states that he has in his collection an old
female silvery-haired bat, taken in New York State, which is brownish
black everywhere, with no trace of the silver tippings. A male in
the Museum of Vertebrate Zoology (no. 13802), taken August 4,
1911, on Kangaroo Creek, Siskiyou County, California, has the mem-
branes and the basal part of the fur everywhere blackish. On the
back, belly, and interfemoral membrane the hairs are tipped with
silvery white. On the face, crown, throat, and a patch over each
shoulder the silvery tips are lacking. This specimen, as evidenced
by the thin, papery skull and the prominent joints of the fingers, is
a juvenal. Merriam (1884, p. 191) states that the young alone possess
the perfect silvery tips to the hairs and that even before going into
winter quarters for the first time their pelage has assumed the grizzled
appearance which characterizes the coats of the adults.
Skull and Teeth—The skull and teeth have been sufficiently de-
scribed under the characterization of the genus (see p. 299).
Average Measurements.—A series of 12 specimens of the silvery-
haired bat, including both sexes, averages in millimeters as follows:
total length, 102.0 (extremes, 92.0-107.0) ; tail vertebrae, 41.1 (34.0—
44.0); tibia, 15.8 (14.0-18.0); foot, 10.0 (9.5-10.0) ; forearm, 39.2
(37.3-42.0) ; greatest leneth of cranium, 16.1 (15.5-16.5).
Synonymy and History.—The silvery-haired bat was first de-
scribed by Le Conte (1831, p. 31) from a specimen from the ‘‘ Kastern
United States’? under the name V{[espertilio|. noctivagans. Peters
(1865, p. 648) made this bat the type species of his new genus
Lasionycteris.
Distribution.—Miller (1912, p. 60) gives the general range of
Lasionycteris noctivagans as ‘‘North America north of Mexico, from
the Atlantic to the Pacific; probably not breeding south of the
Transition Zone.’’ The range in California lies altogether within
the northwestern portion of the state, and for the most part within
the Transition Zone. (See map, text-fig. O.)
302 University of California Publications in Zoology [Vou.17
MEASUREMENTS IN MILLIMETERS OF TWELVE SPECIMENS OF LASIONYCTERIS
NOCTIVAGANS (LECONTE), FROM CALIFORNIA
E SE Se 38
5 a 45° 85) Se
Mus ef ae = i) PEE Se ce
no. Sex se S 2 Gq & 5 2 Nim a7
3281-7 gi 102.0 41.0 17.0 10:05 3920 ees
3282+ 99.0 38.0 17.0 LOO 42:0 ee Paw
184937 gf? 92.0 OO) eee 9.5 15.5 9.3 4.0
3279° 9° 98.0 41.0 15.0 1010305540!) ee at see ac
3280® 9° 106.0 42.0 18.0 10.0 16.4 aie 8.5 4.3
24208 9° 104.0 41°00) ye 8.0 16.2 9.5 7.9 4.2
24212* 9 106.0 42.0 15.8 9.0 16.3 a 8.0 4.2
DAZ NOM LOI eee 44.0 16.0 920) 16.2 era 79 4.1
24211* 9 107.0 42.0 14.3 9.0 aiid?) ESS 9.8 8.0 4.1
24207* © 105.0 42.0 14.7 8.0 39.7 16.5 9.9 8.0 4.2
24209 9 98.0 4010) Bas 8.0 38.4 16.2 9.7 8.0 4.3
24213* 9 105.0 44.0 15.0 9.0 38.3 16.1 = 7.9 4.4
1 From 7000 ft., Mt. Shasta, Siskiyou County.
?From 4 miles north of Oroville, Butte County.
*From McCloud River, near Baird Station, Shasta County.
4From Fyffe, 3700 ft. alt., Eldorado County.
Specimens Examined—The writer has examined 40 specimens of
the silvery-haired bat, from the following localities in California:
Siskiyou County: Kangaroo Creek, 2; Mt. Shasta (at 7000 feet alti-
tude), 2; Shasta County: McCloud River, 15 miles east of Baird
Station, 2; 13 miles east of Baird Station, 2 (Univ. Calif. Dept.
Zool.) ; near Baird Station, 1; Humboldt County: Redwood Creek, 1
(U. S. Nation. Mus.) ; Trinity County: Cafion Creek, 2 (U.S. Biol.
Sury.); Butte County: 4 miles north of Oroville, 1; Mendocino
County: Mt. Sanhedrin, 5 (Acad. Nat. Sci. Phila.) ; Nevada County:
Nevada City, 1 (U. S. Biol. Surv.) ; Eldorado County: Fyffe, 11;
Bijou, 1 (Acad. Nat. Sci. Phila.) ; Marin County: Olema, 1 (Calif.
Acad. Sei.) ; Nicasio, 6 (Amer. Mus. Nat. Hist., 3; U. S. Nation. Mus.,
3); Monterey County: Pacifie Grove, 1 (U. S. Biol. Surv.).
Natural History—The most extended account of the habits of the
silvery-haired bat is that of Merriam (1884, p. 190). His observa-
tions were made in the Adirondack region of New York. Here he
found the bats showing when abroad in the evening a decided liking
for waterways, in some places keeping directly over the water. Sey-
eral bats which were shot and fell into the water swam swiftly and
powerfully through the strong current to the shore, fifteen or twenty
feet distant. A scarcely less favored haunt was the edges of hard-
wood groves, where the bats darted in and out among the branches
in search of insects.
1918] Grinnell: A Synopsis of the Bats of California 303
The flight of this species is described by Merriam as neither so
rapid nor so irregular as that of the red or of the hoary bat.
In the region where Merriam made his observations the young
were born early in July, and were either one or two in number.
MYOTIS ORINOMUS
EUDERMA MACULATUM ili
MYOTIS OCCULTUS
|
LASIONYCTERIS NOCTIVAGANS
SSS
NYCTINOMUS FEMOROSACCUS
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CAI. FORNIA,
Fig. O. Map showing stations of occurrence in California of Myotis orinomus,
Euderma maculatum, Myotis occultus, Lasionycteris noctivagans, and Nyctinomus
femorosaccus, as established by specimens examined by the author.
In California the writer is aware only of summer and fall records
of this bat. The earliest capture is of a female (no. 159941, U.S.
532’
Amer. Mus. Nat. Hist.), at Nicasio, October 19. The species is known
Biol. Sury.), at Pacific Grove, May 25; the latest, of a male (no. ale
304 University of California Publications in Zoology [Vou.17
to be migratory in at least the eastern portion of its range, and the
same probably holds true here in California. The known facts relat-
ing to its migration are recorded on page 232 of the present paper.
Of nine specimens obtained by Joseph Dixon at Fyffe (3700 feet
altitude), Eldorado County, between July 19 and 31, seven were adult
females weighing from 10.2 grams to 12.1 grams each, one was an
immature female weighing 6.3 grams, and one a male, probably imma-
ture, weighing 8.5 grams. Some of the adult females showed evi-
dences of having recently nursed young.
Genus Pipistrellus Kaup
This genus ranges in the Eastern Hemisphere from the northern
to the southern limits of tree growth, and from Tasmania to [reiand.
In the Western Hemisphere it occurs from the northern United States
(except in the Boreal zone’) to southern Mexico (Barrett-Hamilton,
1910, p. 102). The genus contains about forty species, only one of
which occurs in the area under consideration, where it is represented
by two subspecies.
o BY ll 2=2 3-3
Characters —Dental formula: 1 329’ © jay’ PMocg) M 3-3 — 34.
Upper inner incisor about one-third higher than outer one. Lower
incisors all of about equal size and with columnar shafts, separated
from each other by spaces about one-third the width of the shaft;
the abruptly wider crowns are imbricated and deeply trilobed.
Upper canine conical and about twice height of lower canine, which
is similar in form but with a more highly developed cingulum. First
upper premolar minute and lying on inner side of tooth row, crowded
in angle between canine and second upper premolar. Second upper
premolar about midway in height between canine and first molar.
First lower premolar about half height of second and closely crowded
against cingulum of canine; cingulum of this premolar about equal
in width to that of canine. Second lower premolar nearly equal
in height to lower canine. Molar teeth normal for vespertilionid
bats.
Greatest width of skull about two-thirds of the total length of
same. Brain-case distinctly flattened above and forming only a sight
angle with rostrum. Auditory bullae large, their greatest diameter
being distinctly greater than the width of the space between them.
Zygomata slightly expanded. (See pl. 21, fig. 27, and pl. 22,
fig. 39.)
Ear in this genus distinetly longer than broad and tapering to a
narrowly rounded tip. Tragus broadest below its center, its tip
straight or shghtly eurved forwards. Dorsal surface of interfemoral
membrane sprinkled with hairs on basal third. Mammae two.
1918 | Grinnell: A Synopsis of the Bats of California 305
Pipistrellus hesperus hesperus (H. Allen)
Western Bat
Scotophilus hesperus H. Allen (1864, pp. 43-44, figs. 38-40), part. Orig
inal description; type locality, Fort Yuma, California [first locality
mentioned in list of specimens].
Scotophilus hesperus, Cooper (1868, p. 5), part. Distribution.
S[cotophilus]. hesperus, Cooper (in Cronise, 1868, p. 442). Nominal.
Vespertilio (Vesperugo) hesperus, Coues and Yarrow (1875, pp. 94-95).
Diagnosis; distribution.
Vesperugo hesperus, True (1887, p. 515). Concerning the status of V.
hesperus and of V. merriami.
Vesperugo hesperus, Bryant (1891a, p. 358). Nominal.
Vesperugo hesperus, H. Allen (1894, pp. 128-131, pls. 20-21). Deserip-
tion; distribution.
Pipistrellus hesperus, Miller (1897b, pp. 88-90, figs. 20-23), part. Descrip-
tion; distribution.
Pipistrellus hesperus, Elliot (1901, pp. 408-409, fig. 86), part. Diagnosis;
distribution.
Pipistrellus hesperus, Miller and Rehn (1901, p. 259). Type locality.
Pipistrellus hesperus, Elliot (1904a, p. 320), part. Localities of capture
in California.
Pipistrellus hesperus, Elliot (1904b, pp. 582-583, figs. 86, 108), part.
Diagnosis; distribution.
Pipistrellus hesperus, Elliot (1905, pp. 480-481), part. Geographical dis-
tribution.
Pipistrellus hesperus, Stephens (1906, pp. 269-270), part. Description;
distribution; habits.
Pipistrellus hesperus, Elliot (1907, pp. 507-508), part. Localities of cap-
ture in California.
Pipistrellus hesperus; J. Grinnell (1908, pp. 159-160). Distribution and
habits in the San Bernardino Mountains.
Pipistrellus hesperus, Lyon and Osgood (1909, p. 274). Nature and loca-
tion of type.
Pipistrellus hesperus, Miller (1912, p. 60), part. General range.
Pipistrellus hesperus hesperus, J. Grinnell (1913), p. 279), part. Range
in California.
Pipistrellus hesperus hesperus, J. Grinnell (1914, pp. 267-268). Occurrence
on the Colorado River; habits; status of subspecies.
Diagnosis—Size very small (total leneth, 62 to 80 millimeters;
forearm, 26.6 to 30.7 millimeters); tragus blunt, with tip bent
forward; general color of fur buffy gray above, whitish beneath;
membranes, ears, and feet, blackish.
Description: Head.—Muzzle short and broad, the greatest width
of the rostrum equaling its length. Nostrils cireular and directed
outward and downward; region between them shghtly concave. Eye
small, inconspicuous and situated above and posterior to angle of
mouth. A glandular swelling on each side of head between nostril
and eye. Ear short, in an alcoholic specimen reaching only to a
306 University of California Publications in Zoology (Vou. 17
point midway between eye and nostril, when laid forward; anterior
border of auricle strongly convex from the well developed basal notch
to about middle, where it becomes straight and continues so almost
to the broadly rounded tip; posterior border concave immediately
below tip, then strongly convex to basal notch; basal lobe well devel-
oped, separated from main auricle by a deep notch, and joining face
at a point slightly below and behind angle of mouth. Tragus less
than half height of ear, slightly blunt at tip, and curved forward
(text-fig. L). The western bat and the Merriam bat are the only
small bats of California having the tragus curved, and this feature
is sufficient to distinguish them at once from the several small species
of Myotis.
Limbs and Membranes.—Wing short and broad, length of fifth
metacarpal almost equaling that of third. Wing membrane attached
at base of outer toe. Calear very slightly lobed at tip. Tip of tail
projecting somewhat beyond edge of interfemoral membrane. Foot
small, tibia short; length of former scarcely less than half that of
latter. Tail about equal in length to forearm.
Pelage-—Fur everywhere full and soft, 3 to 4 millimeters in length
on body. Dorsal surface of ear furred only on its basal third; a
seanty sprinkling of fine hairs over whole ventral (inner) surface
of ear. Wing membranes naked both above and below, save for a
very narrow strip of fur, about one millimeter in width, which extends
out from side of body. Basal third of interfemoral membrane
sprinkled with fine hairs, both above and below. Toes scantily
clothed, both above and below, with very short, fine hairs.
Color—Hairs everywhere plumbeous-black at bases. Distal two-
thirds of fur on upper surface light buff, brightest on top of head;
on lower surface a very pale tint of light buff, almost white; pelage
everywhere given a grayish cast by the showing through of the darker
bases of the hairs. Ear, muzzle, and wing and tail membranes,
blackish, save that wing membrane is narrowly bordered with pale
eray between foot and fifth finger.
Skull—Form and general characteristics as for the genus. In
general, as has been suggested, the skull of this bat reminds one of
a miniature Lasionycteris.
Measurements—Average and extreme measurements in milli-
meters of a series of twenty western bats, from the Colorado River
Valley, are as follows: ten males: total length, 66.4 (extremes, 62.0—
72.0) ; tail vertebrae, 26.5 (24.0-30.0) ; tibia, 10.6 (10.0-11.5) ; foot,
5.0; forearm, 27.8 (26.6-30.0); greatest leneth of cranium, 11.5
(11.3-11.9). Ten females: total length, 72.9 (69.0-80.0) ; tail verte-
brae, 29.9 (26.0-33.0) ; tibia, 11.2 (10.0-12.0); foot, 5.2 (5.0-6.0) ;
forearm, 29.8 (27.3-30.7); greatest length of cranium, 11.9 (11.7-
112.3).
A comparison of the above averages will show that the females are
as a rule somewhat larger than the males.
Synonymy and History.—The western bat was described by H.
Allen (1864, pp. 48-44) under the name Scotophilus hesperus from
material obtained at Fort Yuma, California, and ‘‘Posa Creek.’’
According to True (1887, p. 515), Dobson in his catalogue of the
307
A Synopsis of the Bats of California
Grinnell
1918]
Mus.
no.
10386
10388
10389
103890
10391
10892
10393
10397
10402
10403
10400
10401
10387
10396
10406
10407
10411
6936
6680
17794
MEASUREMENTS IN MILLIMETERS OF TWENTY ADULT SPECIMENS OF PIPISTRELLUS HESPERUS HESPERUS (H. ALLEN ),
3eX
+O +0 +0 +0 +0 +O +0 +0 40 40 24 OC, OC, BA QA QA OV AOL QA
FROM CALIFORNIA
Locality
Opposite The Needles, Colorado River
Opposite The Needles, Colorado River
Opposite The Needles, Colorado River
Opposite The Needles, Colorado River
Opposite The Needles, Colorado River
Opposite The Needles, Colorado River
Opposite The Needles, Colorado River
20 miles north of Picacho, Colorado River
Potholes, Colorado River
Potholes, Colorado River
8 miles east of Picacho, Colorado River
8 miles east of Picacho, Colorado River
Opposite The Needles, Colorado River
20 miles north of Picacho, Colorado River
Near Pilot Knob, Colorado River
Near Pilot Knob, Colorado River
Near Pilot Knob, Colorado River
Cushenbury Springs, San Bernardino Mts.
Cushenbury Springs, San Bernardino Mts.
Lone Pine Creek, Inyo Co.
Tibia
10.3
10.8
10.0
10.0
10.0
11.0
11.5
11.0
10.8
10.7
12.0
12.0
10.0
11.5
11.0
10.8
11.0
10.8
11.5
11.4
Foot
oy St
oooo
a
AON
Oo
Si
Saoeo se
Or
Forearm
+ Greatest
length of
eranium
a
i)
~ Zygomatic
breadth
>
& Breadth of
brain-case
aan
oono
6.0
Interorbital
constriction
2
—
308 University of California Publications in Zoology (Vou. 17
Chiroptera regards V. hesperus as identical with V. abramus, an Old
World species. Later, Dobson (1886, p. 124) described a new species
of North American Vesperugo under the name of V. merriami, basing
his description on a single specimen sent him by C. H. Merriam.
Dobson gives the type locality as Locust Grove, New York, the home
of Dr. Merriam. Miller (1897b, p. 31) points out the error and
states that the specimen really came from Red Bluff, Tehama County,
California, and places the name merrianv in the synonymy of hes-
perus. J. Grinnell (1913b, pp. 279-280) revives the name merriami
and applies it to the race of hesperus occupying the Lower and Upper
Sonoran zones in California west of the desert divides, thus restrict-
ing the subspecific application of the name hesperus to the desert race.
Distribution—Specimens available indicate that the range of
Pipistrellus h. hesperus oceupies the Lower Sonoran zone chiefly east
of the Pacific divides, comprising the Colorado and Mohave deserts,
and extends from the Mexican line north at least to Coleville, Mono
County (see map, text-fig. P). J. Grinnell (1913), p. 279) gives the
range of this bat as extending west to Fort Tejon, Kern County; but
the present writer finds upon examination of the specimens upon
which this record was based that while Tejon specimens are not typical
merrianu they bear a closer resemblance to that form than to Pipi-
strellus h. hesperus.
Specimens Examined.—The writer has examined 157 specimens
of Pipistrellus hesperus hesperus, from the following localities in
California: Imperial County: Colorado River near Pilot Knob, 18;
Potholes, 4; four miles south of Potholes, 1; eight miles east of
Picacho, 7; twenty miles above Picacho, 7; Thermal, 2; San Diego
County: Carrizo Creek, 3; Santa Ysabel, 14 (U. S. Nat. Mus.) ;
Jacumba, 2 (U. S. Nat. Mus.) ; Vallecito, 4; Dulzura, 6 (Amer. Mus.
Nat. Hist.); Palmetto Spring, 3; Riverside County: Dos Palmos
Spring, Santa Rosa Mountains, 5; Cabezon, 1; Banning, 1; Palm
Springs, 4 (U.S. Biol. Surv., 3; Mus. Vert. Zool., 1) ; Palm Canon, 3;
San Bernardino County: Borax Flat, 3 (U.S. Biol. Surv.) ; Needles,
4 (U. S. Biol. Sury.) ; Colorado River opposite The Needles, 13; east
base Turtle Mountains, 1; Barstow, 1; Victorville, 1; Warren’s
Ranch, 1 (U.S. Biol. Surv.) ; Cushenbury Springs, 5; Kern County:
west slope Walker Pass, 3; Weldon, 1; Onyx, 3; Fay Creek, 3; South
Fork Kern River, 3 (U. S. Biol. Surv.) ; Kern River, 12 miles below
Bodfish, 1; Inyo County: Lone Pine, 4 (Mus. Comp. Zool., 1; U.S.
Biol. Surv., 3); Lone Pine Creek, 5; Independence, 1 (U. 8S. Biol.
1918] Grinnell: A Synopsis of the Bats of California 309
Surv.) ; Death Valley, 4 (U.S. Biol. Surv.) ; Saline Valley, 1 (U.S.
Biol. Surv.); Panamint Valley, 6 (U. S. Biol. Surv.) ; Panamint
Mountains, 5 (U. S. Biol. Surv.) ; Funeral Mountains, 1 (U. 8. Biol.
Surv.) ; Mono County: Coleville, 2 (Mus. Comp. Zool.).
@ PIPISTRELLUS HESPERUS MERRIAMI
& PIPISTRELLUS HESPERUS HESPERUS
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA,
Fig. P. Map showing stations of occurrence in California of Pipistrellus
hesperus hesperus and Pipistrellus hesperus merriami, as established by specimens
examined by the author,
Natural History.—Stephens (1906, p. 270) records these bats as
appearing early in the evening, sometimes soon after sunset, but
states that because of their swift and erratic flight they are difficult
to secure by shooting.
310 University of California Publications in Zoology [Vou. 17
At Cushenbury Springs, on the desert slope of the San Bernardino
Mountains, J. Grinnell (1908, pp. 159-160) found these bats common
in the early part of August. They appeared at late dusk among the
cottonwoods and over the pasture, and swarms of them were to be seen
darting about over the surface of a small pond, frequently dipping
down and touching the surface of the water as if drinking. The five
specimens secured at that time and place were very fat.
In 1908, Grinnell and Swarth (1913, p. 382) found these bats
abundant along Palm Canon and in the neighborhood of Dos Palmos
Spring, on the desert slope of the San Jacinto Mountains. Here they
were found emerging from the crevices of rocks at early dusk. This
was the only species of bat seen abroad after sunrise and before
sunset; in fact one individual was seen in flight about 9 a.m. in the
bright glare of the forenoon sun.
In the spring of 1910 J. Grinnell (1914, pp. 267-268) found the
western bat abundant along the Colorado River between Needles and
Pilot Knob. Here it was noticed that the bats were to be found only
in the neighborhood of cliffs and rocky hillsides.
No instances of the occurrence of this bat in or about buildings
or as hanging up on the foliage of trees, have been recorded.
Specimens in the Museum of Vertebrate Zoology have been taken
in all the months from February to August, inclusive, and there is
also a specimen taken at Palm Springs in December. Winter collect-
ing would probably reveal the presence of this species throughout
the year in at least the lowest parts of its range. Stephens (1906,
pp. 269-270) states that very few western bats remain in California
in winter. ‘‘The northward migration is at its height about the end
of March, at which time’’ the bats are abundant about certain springs
along the western border of the Colorado Desert.
The number of young and time of breeding are indicated by the
following data, taken from specimens in the Museum of Vertebrate
Zoology secured in Kern County in the summer of 1911: no. 16308,
taken June 18, contained two embryos; no. 16309, taken June 19,
contained two embryos; no. 16310, taken June 20, contained two em-
bryos; no. 16312, taken June 23, contained two embryos; no. 16313,
taken July 1, contained two embryos; no. 16314, taken July 11,
contained one embryo.
a aa
1918] Grinnell: A Synopsis of the Bats of California 311
Pipistrellus hesperus merriami (Dobson)
Merriam Bat
Scotophilus hesperus H. Allen (1864, pp. 43-44, figs. 38-40), part. Orig-
inal description of hesperus; Posa Creek [Kern County] specimens
referable to merriamt.
Scotophilus hesperus, Cooper (1868, p. 5), part. Distribution.
Vesperugo merriami Dobson (1886, p. 124). Original description; type
stated to be from Locust Grove, New York, but really from Red Bluff,
Tehama County, California.
Vesperugo merriami, True (1887, p. 515). Validity doubted.
Vesperugo merriami, Bryant (1892, p. 223). Nominal.
Vesperugo merriami, Miller (1897b, p. 31). Nomenclature.
Pipistrellus hesperus, Miller (1897b, pp. 88-90), part. Description; gen-
eral distribution.
Pipistrellus hesperus, Elliot (1901, pp. 408-409, fig. 86), part. Diagnosis;
general distribution.
Pipistrellus hesperus, Elliot (1904b, pp. 582-5838, figs. 108, 86), part.
Diagnosis; general distribution.
Pipistrellus hesperus, Stone (1904a, p. 579). Reeord of occurrence in
Mendocino County.
Pipistrellus hesperus, Elliot (1905, pp. 480-481), part. General distri-
bution.
Pipistrellus hesperus, Stephens (1906, pp. 269-270), part. Description ;
distribution; habits.
Pipistrellus hesperus, Elliot (1907, pp. 507-508), part. Loealities of cap-
ture in California.
Pipistrellus hesperus, Miller (1912, p. 60), part. General range.
Pipistrellus hesperus merriani, J. Grinnell (1913b, pp. 279-280). Range
in California.
Pipistrellus hesperus merriami, J. Grinnell (1914, pp. 267-268). Status
and range.
lo
Diagnosis—Size very small (total length 66 to 78 millimeters,
forearm 27.5 to 30.8); tragus blunt, with tip bent forward; general
color of fur buffy brown both above and below; membranes, ears, and
feet, blackish.
Description.—This subspecies, Pipistrellus hesperus merriami, very
closely resembles the preceding one, Pipistrellus hesperus hesperus,
differing only in darker coloration and in slightly larger average size.
Color—Wing and tail membranes, ears and feet, blackish (see
pl. 18, fig. 16). Terminal portion of body fur, both above and
beneath, nearest ‘‘warm buff’’ of Ridgway’s Color Standards; but
the darker bases of the hairs showing through lend a brownish aspect
to the mass appearance.
Measurements—Average and extreme measurements in milli-
meters of a series of 16 specimens of the Merriam bat in the Museum
of Vertebrate Zoology are as follows: six males: total length, 69.0
(extremes, 66.0—-73.0) ; tail vertebrae, 29.0 (28.0-30.0) ; tibia, 11.0
(10.6-12.0) ; foot, 5.5 (5.0-6.5) ; forearm, 28.1 (27.5-29.0) ; greatest
312 University of California Publications in Zoology [Vou.17
length of cranium, 11.6 (11.38-11.8); ten females: total length, 73.6
(67.0-78.0) ; tail vertebrae, 29.2 (26.0-34.0) ; tibia, 11.4 (11.1-11.7) ;
foot, 5.6 (5.0-6.0); forearm, 29.6 (28.0-30.8); greatest length of
eranium, 12.0 (11.9-12.2).
Synonymy and History—The synonymy and history of this bat
is ineluded with that of P. h. hesperus on page 306. In his original
description of Pipistrellus hespcrus, H. Allen (1864, pp. 43-44) does
not designate a type, but lists three specimens, the first being from
Fort Yuma and the two remaining from Poso Creek (Kern County).
Miller (18970, p. 88) designates the Fort Yuma specimen (no. 5406
U. S. Nation. Mus.) as the type of hesperus. I have examined the
two skins (nos. 5509 and 5570 U. S. Nation. Mus.) from Poso Creek
and find them to be Pipistrellus hesperus merriami, as would be ex-
pected from the geographic position of this locality.
Distribution—Specimens examined indicate the range of this bat
as lying within the Upper Sonoran and Transition zones altogether
west of the desert divides, from the Mexican line northwest through
the San Diegan district and through the San Joaquin and Sacramento
valleys, east of the humid coast belt and west of the Sierra Nevada, to
Butte and Tehama counties (see map, text-fig. P). Specimens from
Witch Creek (or Santa Ysabel), San Diego County, are allocated with
difficulty, some appearing to be nearest hesperus and others nearest
merriamt. .
Specimens Examined.—The writer has examined 46 specimens
of Pipistrellus h. merriami, from the following localities in California :
San Diego County: Twin Oaks, 2 (U. S. Biol. Surv.) ; Escondido, 1;
Witch Creek, 4 (San Diego Soe. Nat. Hist., 1; U.S. Nation. Mus., 3) ;
Riverside County: San Jacinto Mountains, 2 (U. 8. Nation. Mus.) ;
San Jacinto Lake, 5 (U. S. Nation. Mus.); Los Angeles County:
Pasadena, 1; Rubio Wash, near Pasadena, 3; San Francisquito Canon,
1; San Gabriel Cafion, 2 (U. S. Biol. Surv.) ; Kern County: Grape-
vine Canon, near Fort Tejon, 3; San Emigdio, 1 (U. 8. Biol. Surv.) ;
Poso Creek, 2 (U. S. Nation. Mus.); San Luis Obispo County:
Cuyama Valley, 1; Mariposa County: El Portal, 1; Pleasant Valley,
2; Yosemite Valley, 3; Madera County: Raymond, 3; Solano County:
three miles west of Vacaville, 1; Yolo County: Rumsey, 3; Eldorado
County: Limekiln, 2; Fyffe, 1; Sutter County: Butte Slough, 1; Men-
docino County: Mt. Sanhedrin, 1 (Acad. Nat. Sei. Phila.).
Natural History—tThe time of breeding and number of young is
indicated by the following data, taken from specimens in the Museum
of Vertebrate Zoology : female, no. 18497, taken near Vacaville, Solano
a
f the Bats of Californ
A Synopsis o
Grinnell
1918]
MEASUREMEN'TS IN MILLIMETERS OF SIXTEEN
Mus.
no.
6937
5161
16669
14648
14646
18494
6684
6681
6683
6682
5162
5164
14647
18497
18496
18495
+0 +0 +0 +0 40 40 40 10 1010 0,0,0,0,0, 0,
Locality
Rubio Wash, near Pasadena, Los Angeles Co.
Grapevine Canon, near Ft. Tejon, Kern Co,
Cuyama Valley, San Luis Obispo Co.
Raymond, Madera Co.
Raymond, Madera Co.
Butte Slough, 1 mi. west Butte, Sutter Co.
San Francisquito Cafion, Los Angeles Co.
Grapevine Canon, near Ft. Tejon, Kern Co.
Rubio Wash, nr. Pasadena, Los Angeles Co.
Pasadena, Los Angeles Co.
Rubio Wash, nr. Pasadena, Los Angeles Co.
Escondido, San Diego Co.
Raymond, Madera Co.
3 mi. west Vacaville, Solano Co.
Rumsey, Yolo Co.
Rumsey, Yolo Co.
ADULT SPECIMENS OF PIPISTRELLUS
Foot
yas
a)
2s
cor)
al
ooo nooo co oe
AS
co © Forearm
re)
bo bo
27.5
bo
oe
So
po
4
Nn
29.0
30.0
29.0
28.0
30.6
30.6
29.0
30.0
30.8
28.8
length of
cranium
oo oO
me » Greatest
ee
~a ~ Zygomatic
ti “breadth
7.3
6.8
7.7
8.0
8.0
7.8
8.0
7.7
7.5
8.0
7.8
brain-case
i» to
i) Ge
S tw
co o Interorbital
HESPERUS MERRIAMI (DOBSON), FROM CALIFORNIA
constriction
te
3.5
3.4
314 University of California Publications in Zoology [Vou.17
County, July 2, 1912, two embryos; female, no. 18495; taken at Rum-
sey, Yolo County, June 24, 1912, two embryos. Two examples of the
Merriam bat secured by Dixon at Limekiln, Eldorado County, August
2, 1916, weighed as follows: no. 24216, female, 4 grams; no. 24217,
male, 3.5 grams.
Genus Eptesicus Rafinesque
According to Miller (1907, p. 208) this genus is represented in
Africa, Madagascar, Australia, Asia (except the Malay region), and
America from southern Canada southward (except the Lesser An-
tilles). About forty-five species belonging to this genus are known.
In North America only one species, consisting of several races, is
known to exist. Among the species of the Vespertilionine group
oceurring in California the one belonging to the genus Eptesicus may
be readily distinguished by its comparatively large size (total length,
105 to 122 millimeters), reduced dental formula, and nearly uniform
brown coloration of the body.
99) — = 2!
Characters ——Dental formula: 1 —— sl II Se
5-9) iar PM g=o) Masa
Upper incisors both well developed, the outer one much the smaller,
reaching barely to cingulum of inner one. Lower incisors almost
uniform in size, forming a closely crowded convex row between the
canines, with their crowns overlapping; the crowns tri-lobed, and that
of the outermost slightly the largest. Canines, both above and below
simple, each with distinct cingulum but no secondary cusp. Height
of single upper premolar exceeds that of any molar. First lower
premolar about half the size of the second.
Skull flattened above; angle between brain-case and rostrum
slight; sagittal crest conspicuous. (See pl. 23, fig. 52; pl. 24, fig. 60).
Ears short (17 to 20 millimeters in height from meatus) and some-
what narrower than long; basal lobe well developed but not excessively
large, being in length about half height of tragus. Tragus straight,
short (7 to 9 millimeters in height), and directed slightly forward,
broadest near middle, and tapering to a slightly blunted point. Mem-
branes nearly naked. Mammae two.
Eptesicus fuscus (Peale and Beauvois)
Large Brown Bat
Vespertila fuscus Peale and Beauvois (1796, p. 14). Original description ;
type locality, Philadelphia.
Scotophilus fuscus, H. Allen (1864, pp. 31-35, figs. 27-29). Deseription;
nomenclature; specimens listed from San Francisco and Posa Creek.
Scotophilus fuscus, Cooper (1868, p. 5). Distribution.
1918]
Grinnell: A Synopsis of the Bats of California 315
Scotophilus fuscus, Cooper (in Cronise, 1868, p. 442). Occurrence in Cali-
fornia.
Vesperugo serotinus var. B, Dobson (1878, pp. 191-194). Description;
specimen listed from Monterey.
Vesperugo serotinus, C. H. Townsend (1887, p. 182). Record of two
specimens secured at northeast base of Mt. Shasta, Siskiyou County.
Adelonycteris fuscus, H. Allen (1894, pp. 112-121, pls. 15-17). Deserip-
tion; habits; specimens listed from Santa Barbara and San Francisco.
Vespertilio fuscus, Miller (1897b, pp. 96-99, figs. 24-26). Description;
distribution; many California localities.
Vespertilio fuscus, Merriam (1899, p. 87). Occurrence on Mt. Shasta.
Vespertilio fuscus, Elliot (1901, p. 410). Diagnosis; distribution.
Eptesicus fuscus bernardinus Rhoads (1901, pp. 618-619). Original
description of supposed new form from San Bernardino Valley; dis-
tribution.
Vespertilio fuscus bernardinus, Miller and Rehn (1903, p. 122). Type
locality.
Vespertilio fuscus, Elliot (1904a, p. 320). Record stations in California.
Eptesicus fuscus bernardinus, Elliot (1904b, p. 589). Distribution; gen-
eral characters.
Eptesicus fuscus, Stone (1904a, p. 579). Occurrence at Sanhedrin Moun-
tain, Mendocino County.
Eptesicus fuscus melanopterus Rehn (in Stone, 1904b, pp. 590-591). Orig-
inal description of supposed new form from Mt. Tallac, Eldorado
County.
Vespertilio fuscus melanopterus, Elliot (1905, p. 483). Distribution; gen-
eral characters.
Vespertilio fuscus bernardinus, Elliot (1905, p. 483). Distribution; gen-
eral characters.
Eptesicus fuscus bernardinus, Stephens (1906, p. 270). Diagnosis; dis-
tribution.
Eptesicus fuscus melanopterus, Stephens (1906, pp. 270-271). Diagnosis;
distribution.
Vespertilio fuscus, Elliot (1907, pp. 509-510). Localities of capture in
California: Panamint Mountains; Fort Tejon; Mount Whitney.
Vespertilio fuscus, J. Grinnell (1908, p. 159). Occurrence in the San
Bernardino Mountains.
Vespertilio fuscus bernardinus, J. Grinnell (1908, p. 159). Discussion
of validity of subspecies.
Eptesicus fuscus, Seton (1909, p. 1179). Map showing actual record
stations and hypothetical range.
[Eptesicus fuscus| bernardinus, Seton (1909, p. 1179). Map showing
actual record stations and hypothetical range.
[Eptesicus fuscus] melanopterus, Seton (1909, p. 1179). Map showing
actual record stations and hypothetical range.
Eptesicus fuscus melanopterus, Miller (1912, p. 62). - Type locality.
Eptesicus fuscus bernardinus, Miller (1912, p. 62). Type locality.
Eptesicus fuscus fuscus, J. Grinnell (1913b, p. 280). Range in California.
Eptesicus fuscus fuscus, Grinnell and Swarth (1913, pp. 381-382). As
occurring in the San Jacinto Mountains.
Eptesicus fuscus, J. Grinnell (1914, p. 268). As occurring along the
Colorado River.
316 University of California Publications in Zoology [Vou.17
Diagnosis.—Size medium (total length 107 to 122 millimeters) ; .
ear about 18 millimeters in height, its width slightly less than two-
thirds of height; tragus two-thirds height of ear, tapermg, and
directed slightly forwards. Color, varying dorsally from raw umber
to vandyke brown, ventrally from pale wood brown to light bister.
Description: Head.—Muzzle short and broad, greatest width of
rostrum almost equaling its length. Nostrils elliptical in shape and
directed outward and slightly downward; region between them
slightly coneave. Hye small and inconspicuous and situated above
posterior angle of mouth. A tumid, glandular area, some three or
four millimeters in width, occupies entire space between nostril and
eye on each side of head and adds to apparent width of rostrum, Ear
short, reaching barely to nostril when laid forward. (See pl. 18,
fig. 13.)
Limbs and Membranes.—Wing short and broad (text-fig. E),
length of fifth metacarpal almost equaling that of third. Wing
membrane attached to foot a little beyond bases of toes. Free edge
of interfemoral membrane a little shorter than calear and terminating
at base of next to last caudal vertebra. Foot more than half length
of tibia. Calear slightly longer than foot, keeled on its outer edge,
and terminating in a faintly defined lobe.
Pelage—F ur everywhere full and soft; on dorsal surface of body
about 7 millimeters in length, on ventral surface slightly shorter.
Ears furred only on basal third of outer surface; scantily sprinkled
with hairs over most of inner surface, these hairs being most numerous
near anterior border of auricle. Dorsal surface of wing naked, save
for narrow strip of fur continuous with fur of body. Ventral surface
of wing membrane furred in a manner similar to that of dorsal sur-
face. Interfemoral membrane furred only at its extreme base above;
but below, scattering hairs extend almost to tip of tail. Toes scantily
clothed, both above and below, with very fine, short hairs.
Color.—The color varies considerably, skins from even a single
locality exhibiting a wide range. An average condition of coloration
is shown by a brown bat taken at Kenworthy, in the San Jacinto
Mountains, in May, 1908, as follows: above, bases of hairs dark seal
brown, their distal two-thirds raw umber; hairs on throat pale wood
brown; rest of fur-on lower surface pale seal brown on basal half, the
distal half pale wood brown. Ears and membranes blackish.
Skull and Tecth—As deseribed for the genus.
Measurements—Average and extreme measurements in milli-
meters of a series of ten brown bats in the Museum of Vertebrate
Zoology are as follows: five males: total length, 109.8 (extremes
107.0-112.0) ; tail vertebrae, 43.8 (38.0-47.0) ; tibia, 16.9 (16.0-17.5) ;
foot, 9.0 (8.0-10.0); forearm, 44.1 (43.8-44.9); greatest length of
cranium, 18.3 (17.9-18.8) ; five females: total length, 116.1 (110.0-
119.0) ; tail vertebrae, 45.3 (43.0-48.0) ; tibia, 17.3 (16.4-18.0) ; foot,
10.0 (9.0-12.0) ; forearm, 46.2 (42.9-49.0) ; greatest length of skull,
19.0 (18.8-19.4).
A comparison of the above figures will show that the females aver-
age somewhat larger than the males.
ud
{ the Bats of Californ
ynopsis o
A &
Grinnell
1918}
MEASUREMENTS IN MILLIMETERS OF TWENTY-ONE ADULT EXAMPLES OF EPTESICUS FUSCUS (PEALE
Mus.
no.
1864
5173
5177
5179
5176
5141
20813
18499
8904
11840
2246
6688
6697
6942
5144
10697
14649
18498
20814
11841
11842
ive}
o
*
+0 +0 +0 40 40 40 40 1010 1010 E,0Q,Q QQ Q0Q,Q AQ
Locality
Kenworthy, San Jacinto Mts.
S. Fork Santa Ana River, San Bernar-
dino Mts.
Cushenbury Springs, San Bernardino Mts.
Bluff Lake, San Bernardino Mts.
Bluff Lake, San Bernardino Mts.
Mt. Pinos, 6500 ft., Ventura Co.
1 mi. west Guerneville, Sonoma Co.
Rumsey, Yolo Co.
Sherwood, Mendocino Co.
Eureka, Humboldt Co.
Hemet Lake, San Jacinto Mts.
Arroyo Seco, near Pasadena
Arroyo Seco, near Pasadena
Arroyo Seco, near Pasadena
Fort Tejon, Kern Co.
Pilot Knob, Colorado River
Raymond, Madera Co.
Mill Creck, 2 mi. N.E. Tehama, Tehama Co.
1 mi. west Guerneville, Sonoma Co.
Eureka, Humboldt Co.
Yureka, Humboldt Co.
108.0
111.0
112.0
107.0
110.0
112.0
124.0
112.0
112.0
119.0
110.0
118.0
116.0
119.0
107.0
116.0
119.0
120.0
122.0
120.0
vertebrae
i Daal
44.0
38.0
47.0
45.0
46.0
44.0
52.0
45.0
50.0
48.0
43.0
45.0
47.0
45.0
44.0
48.0
46.0
49.0
53.0
49.0
reatest
length of
cranium
uo G
SS
_
17.8
18.4
18.8
18.0
18.5
18.9
19.1
19.2
19.5
19.0
19.0
18.8
19.4
18.8
18.3
19.2
19.2
19.8
20.4
20.0
Zygomatic
breadth
11.8
Breadth of
brain-case
oO
bo
9.5
9.4
9.3
9.4
9.4
10.4
10.0
9.8
MY)
10.0
9.5
10.0
10.1
10.0
9.6
OFT
10.1
10.2
10.2
10.2
AND BEAUVOIS), FROM CALIFORNIA
+ Interorbital
< constriction
f=)
4.7
4.6
318 University of California Publications in Zoology [Vou.17
Synonymy and History—The brown bat was described by Peale
and Beauvois (1796, p. 14) under the name Vespertila (perhaps mis-
print for Vespertilio) fuscus from material collected at Philadelphia,
Pennsylvania.
Eptesicus fuscus bernardinus was described by Rhoads (1901,
p. 619) from a single specimen taken near San Bernardino, California.
‘
This individual is described as ‘‘pallid bistre above, brownish drab
below; .. . wing membranes and ears in bernardinus very dark.”’
Eptesicus fuscus melanopterus was described by Rehn (in Stone,
19046, pp. 590-591) from five specimens obtained at Mount Tallac,
“‘above rather
dark cinnamon, lightest on the top of the head and at the shoulders.
Under surface reddish wood brown. Membranes and face deep
Eldorado County, California. The color is given as
blackish.’’
As explained beyond, under General Remarks, the latter two
names had best be kept in synonymy until better reasons have been
brought forward than have so far been advanced for the recognition
of more than one subspecies within the state.
Distribution.—The range of this species extends over the whole
of the United States and into British Columbia (Miller, 1897), p. 96).
In California it oeeurs chiefly in the Upper Sonoran and Transition
zones, where it has been found practically throughout the state. (See
map, text-fig. Q.).
Specimens Examined.—The writer has examined 278 specimens
of the large brown bat from the following localities in California:
Siskiyou County: Kangaroo Creek, 3; Mt. Shasta, 6 (U. 8. Nation.
Mus., 3; Mus. Vert. Zool., 3); Shasta County: near Baird Station, 4
(Mus. Vert. Zool., 1; U. C. Dept. Zool., 3); Potter Creek Cave, 2
(U. C. Dept. Zool.) ; Humboldt County: Eureka, 8; Mendocino
County: Sherwood, 2; Covelo, 1; Mt. Sanhedrin, 2 (Acad. Nat. Sci.
Phila.) ; Tehama County: Mill Creek, 1; Yolo County: Rumsey, 1;
Sacramento County: Folsom, 1 (Mus. Comp. Zool.) ; Colusa County :
Snow Mountain, 1 (Stanford Univ.) ; Nevada County: Nevada City, 6
(U. S. Biol. Surv.) ; Placer County: Colfax, 8 (Stanford Univ.) ;
Eldorado County: Mt. Tallae, 5 (Acad. Nat. Sci. Phila.) ; Fyffe, 13;
Limekiln, 6; Echo, 1 (Stanford Univ.); Placerville, 3 (Stanford
Univ.) ; Fallen Leaf Lake, 1; Sonoma County: Cloverdale, 1 (U. 8.
Nation. Mus.) ; Cazadero, 1; Guerneville, 3; Marin County: Nieasio,
24 (U. S. Nation. Mus.); Alameda County: Sunol, 1 (U. S. Biol.
Surv.) ; Niles Canon, 2 (Calif. Acad. Sei.) ; San Mateo County: Pes-
1918 | Grinnell: A Synopsis of the Bats of California 319
eadero Creek, 3; Santa Clara County: Mountain View, 1 (Stanford
Univ.) ; Monterey County: Monterey, 5; Pacific Grove; 1 (Stanford
Univ.) ; Mariposa County: Yosemite Valley, 7 (U.S. Nation. Mus., 3;
Mus. Vert. Zool., 4) ; Mereed Lake, 7; Madera County: Raymond, 1;
So
G
ep
!
@ EPTESICUS FUSCUS
oe
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA,
g. Q. Map showing stations of occurrence in California of Eptesicus fuscus,
as SE oined by specimens examined by the author. This species is most wide-
ranging of all Californian bats.
Tulare County: Little Kern River, 3 (U.S. Biol. Surv.) ; Whitney
Meadows, 1; Taylor Meadow, 1; Trout Creek, 6; Kern County: Posa
Creek, 2 (U. 8. Nation. Mus.) ; head of Kelso Valley, 1; Fay Creek, 2;
Kernville, 1 (U. S. Biol. Surv.) ; Walker Basin, 4 (U.S. Biol. Surv.) ;
Kern Lakes, 1 (U. 8. Biol. Surv.) ; Tehachapi, 1 (U. S. Biol. Surv.) ;
320 University of California Publications in Zoology [Vou.17
Fort Tejon, 3; Ventura County: Mount Pinos, 3; San Bernardino
County: Providence Mountains, 2 (U. S. Biol. Surv.) ; San Bernar-
dino Valley, 1 (Acad. Nat. Sci. Phila.) ; San Bernardino Mountains,
31; Los Angeles County: near Pasadena, 14; west fork of San Gabriel
River, 1; Orange County: Trabuco Canon, 1; Riverside County:
Riverside, 1 (Stanford Univ.); San Jacinto, 32 (Stanford Univ.) ;
San Jacinto Mountains, 10; San Diego County: Escondido, 2; Foster,
1; Cuyamaca Mountains, 6; San Felipe Canon, 2; Julian, 5; Pine
Valley, 4 (U. S. Nation. Mus.) ; Dulzura, 2 (U. S. Nation. Mus.) ;
Imperial County: Colorado River near Pilot Knob, 1; Palo Verde, 1.
Natural History. J. Grinnell (1908, p. 159) reports the brown
bat to have been the most common and generally distributed bat in
the San Bernardino Mountains in the summers of 1905, 1906, and
1907. It came out early in the evening, often soon after sundown,
and proved easy to secure by shooting. Specimens taken in August
were excessively fat.
At Guerneville, Sonoma County, where three specimens were
secured in the summer of 1913, these bats were invariably seen flying
slowly and steadily high over the canon bed, whereas the hoary bats
flew low over the meadows, and the little California bats zigzagged
in and out close about the tops of the young redwood growth. The
large size and slow, steady flight of this bat render it comparatively
easy to secure by shooting; hence the abundance of specimens in
collections misrepresents its actual relative abundance among the
various species of bats in the wild.
Bailey (1905, p. 211), when recording the large brown bat from
Texas, states that he found two lower jaws of this bat, among numer-
ous other bones, in pellets under the nest of a great horned owl.
Three females in the Museum of Vertebrate Zoology, each con-
taining a single embryo, were secured on the following dates: April
22, June 10, and June 25. Adult females secured by Dixon at Fyffe,
Eldorado County, between July 19 and 31, were considered by him
(MS) as having ceased to nurse their young.
General Remarks——An examination of a series of 278 specimens
of the large brown bat, taken in California, shows the probable exist-
ence of two or even three races within the state. Specimens from the
humid coast belt of northwestern California average very much
darker than a series from the southeastern part of the state; yet
individual variation is so great that until a much larger series of
specimens becomes available for comparison it is impossible to define
1918] Grinnell: A Synopsis of the Bats of California 821
limits of range satisfactorily. For example, five specimens of EH ptesi-
cus fuscus in the Museum of Vertebrate Zoology, selected because
of their close resemblance in color (bright buffy brown above, pale
avellaneous beneath) were found to have come from the following
widely separated localities: No. 2778, Julian, San Diego County;
no. 6959, San Bernardino Mountains, San Bernardino County; no.
5145, near Pasadena, Los Angeles County; no. 5141, Mt. Pinos,
Ventura County; no. 3286, Mt. Shasta, Siskiyou County.
In the original description of EF. f. bernardinus (from a single
specimen taken in the ‘‘San Bernardino Valley’’), Rhoads (1901,
p- 619) remarks that ‘‘a series of four specimens from the same
collector taken in the ‘San Bernardino Mts.’ in September, 1893,
shows that the mountain form is inseparable from fuscus; one of
these, however, is a perfect intergrade.”’ I have examined Rhoads’
type (no. 8247, ¢, Acad. Nat. Sci. Phila.) and find it well within
the range of individual variation of specimens from the San Bernar-
dino Mountains, and darker than the two specimens at hand from the
Colorado River.
J. Grinnell (1908, p. 159) comments as follows upon a series of
skins of the large brown bat taken in the San Bernardino Mountains:
The series of thirty skins secured shows much variation in depth of color.
Some are very light-colored, and these agree with Rhoads’ subspecies bernardinus.
But others are as dark as the darkest I have seen from elsewhere in central and
southern California, so that I cannot perceive the existence of a race bernardinus
if it is to be based on color characters alone; unless it be that all California
examples differ from the eastern animal.
Grinnell and Swarth (1913, p. 382) remark concerning a series
of nine large brown bats taken in the San Jacinto Mountains: ‘‘The
specimens taken show much variation in color, and the remarks made
in regard to a series from the San Bernardino Mountains apply here.’’
J. Grinnell (1914, p. 268) says of a single example of Eptesicus
fuscus taken by the Colorado River expedition on the California side
of the river near Pilot Knob:
The specimen secured (no. 10697) is an adult female. It appears to differ
in small size and extreme paleness from the average of the species from California.
It about equals in the latter respect the palest out of a series of ninety brown
bats from the Pacific slope of California. The color dorsally is uniform isabella
color, ventrally pale wood brown. Measurements: length 107 mm., tail vertebrae
44, foot 9, forearm 42.5, longest finger 72, ear (dry) 12. The skull, too, is
appreciably small.
A general inspection of the Museum’s series of this species from California
points towards the existence within the state of at least three geographic races
322 University of California Publications in Zoology [Vou.17
based on size and depth of color. But so much of the total area is unrepresented
by specimens that systematic analysis at this time seems inadvisable.
Stone (1904a, p. 579) says of two specimens, male and female, of
Eptesicus fuscus taken at Mount Sanhedrin, California: ‘‘These speci-
mens are identical with topotypes of HZ. fuscus from Philadelphia.”’
Rehn (in Stone, 1904b, p. 590) states that H. f. melanopterus
which he describes from Mount Tallae
.is no doubt closest related to f. osceola than any of the form[s] of fuscus.
The original series of the former has been examined in this connection, and the
differential characters were drawn from it. The relationship with true fuscus
is not so close as an examination of a series of thirty specimens from, or within
a radius of, twenty miles of the type locality shows. No close relationship exists
with E. f. bernardinus Rhoads, which is a very pale type quite different from
any of the forms here considered.
I have examined the type and four paratypes of EF. f. melanop-
terus, which are in the Academy of Sciences of Philadelphia, and find
them very nearly uniform in coloration with, and well within the
range of individual color variation in, series of specimens from else-
where in the United States and Canada. As Rehn states (loc. cit.),
the skull is identical with that of the typical form.
Taking all the above testimony into consideration I feel that the
subspecifie splitting of Hptesicus fuscus in California is a difficult
problem, and one which should not be undertaken until abundant
material is available for comparison, from elsewhere in the United
States and from the adjoining provinces of Canada and Mexico.
Genus Nycteris Borkhausen
This genus ranges from Central America north to the limits of
tree growth in northern North Ameriea. It occurs also on the Greater
Antilles, and on the Bahama; Galapagos and Hawaiian islands (Mil-
ler, 1907, p. 221). Two distinet species are known to occur north of
Panama, one of which is divisible into at least five geographic races.
2 lll a ‘1 ee 3-3
Characters—Dental formula: 1 303" 1 PMo-9> Mg79 = 82.
Upper incisors only two, short and ee ge oa converging at
tips; height from cingulum to tip of crown less than twice greatest
diameter. of tooth. Lower incisors all closely alike, the shafts being
widely separated and columnar in form; crowns abruptly widened,
being at their bases twice width of shafts; crowns trilobed and imbri-
cated; outer pair of lower incisors closely crowded against bases of
canines. Canines well developed, the upper slightly the larger.
1918] Grinnell: A Synopsis of the Bats of California 323
Anterior upper premolar minute, and crowded into inner angle be-
tween canine and large second premolar. First lower premolar
scarcely half height of second. Cusps of lower molars more devel-
oped than those of upper molars.
Skull short and broad, greatest width at least two-thirds its length.
Rostrum about two-thirds length of brain-case, and sloping abruptly
downward from it anteriorly. (See pl. 23, figs. 49, 51; pl. 24, figs.
57, 59.) Auditory bullae greater in diameter than space between
them. Zygomatiec arches not expanded. Ear short and rounded;
when laid forward reaching barely to mouth. Fifth finger much
shorter than third, the difference between them about equaling length
of thumb. Interfemoral membrane very large; most of its upper
surface furred; length of tail vertebrae exceeding that of forearm.
Mammae four.
Nycteris borealis teliotis (11. Allen)
Western Red Bat
Lasiurus noveboracensis, H. Allen (1864, pp. 15-20, figs. 13-17), part.
Description; general distribution.
Lasiurus noveboracensis, Cooper (1868, p. 5). Distribution.
Lasiurus noveboracensis, Cooper (in Cronise, 1868, p. 442). Occurrence
in California.
Atalapha teliotis H. Allen (1891, pp. 5-7). Original description of the
subspecies found in California; precise type locality unknown.
Atalapha noveboracensis, Bryant (1891a, p. 358). Nominal.
Atalapha teliotis, Bryant (1891b, p. 113). Nominal.
Atalapha teliotis, Bryant (1892, p. 219). Nominal.
Atalapha teliotis, H. Allen (1894, pp. 153-155, pls. 27-28). Description.
Lasiurus borealis teliotis, Miller (1897b, pp. 110-111, figs. 29-30). De-
scription; general distribution.
Lasiurus borealis teliotis, Elliot (1901, p. 415). Diagnosis; general dis-
tribution.
Lasiurus borealis teliotis, Miller and Rehn (1901, p. 262). Type locality
(California).
Lasiurus borealis teliotis, Rehn (in Stone, 1904b, p. 591). Record of
capture at Linden, San Joaquin County, June 2, 1898.
Lasiurus borealis teliotis, Elliot (1904b, pp. 593-594). Distribution;
diagnosis.
Lasiurus borealis teliotis, Elliot (1905, p. 486). Distribution.
Lasiurus borealis teliotis, Stephens (1906, p. 271, 1 fig. in text). Deserip-
tion; distribution.
Lasiurus borealis teliotis, Elliot (1907, p. 513). Reeord of specimen
taken at Monterey.
Lasiurus borealis teliotis, Lyon and Osgood (1909, p. 277). Nature and
location of type.
Lasiurus borealis teliotis, Seton (1909, p. 1185). Map showing record
stations and hypothetical range.
Nycteris borealis teliotis, Miller (1912, p. 64). General range.
Nycteris borealis teliotis, J. Grinnell (1913b, p. 280). Range in Cali-
fornia.
324 University of California Publications in Zoology [Vou.17
Diagnosis —Size medium (forearm 34 to 41.8 millimeters long) ;
whole dorsal surface of interfemoral membrane furred; color ranging
from rufous red or fawn to yellowish gray, some of the hairs tipped
with whitish.
Description: Head.—Muzzle short and broad; nostrils directed
outward and downward, apertures in an alcoholic specimen 2 milli-
meters apart. Eye small and inconspicuous. A glandular swelling
on each side of head between nostril and eye. In an alcoholic speci-
men the ears when laid forward reach barely to the mouth. Anterior
border of ear strongly but irregularly convex from free point of
anterior basal lobe to tip; through this portion of its periphery it
forms almost a semicircle. Posterior border of ear concave from tip
to one-third distance to posterior basal lobe, and convex on remaining
two-thirds. Tragus slightly more than half height of ear, and tri-
angular in general outline with tip directed forwards.
Limbs and Membranes——Wing attached at base of toes. Foot
less than half leneth of tibia; claws stronely curved. Calear slender,
about twice as long as foot and considerably shorter than free border
of interfemoral membrane; not lobed at tip in the specimens at
hand. Tail unusually long (5 to 15 millimeters longer than forearm)
and enclosed to extreme tip in interfemoral membrane (pl. 18, fig. 14).
Pelage.—¥ ur everywhere full and soft; longest on dorsal surface
between shoulders, where the maximum length is about 10 millimeters.
Ear furred only on basal two-thirds of its dorsal surface; ventral
surface of ear scantily haired along borders; otherwise naked. Fur
on dorsal surface of wing continuous with that on body to a line
running from ankle jomt to about middle of humerus; otherwise
naked save for three small patches of hair, the first lying at base
of thumb, the second extending along both sides of basal third of
fifth metacarpal, and the third occupying anterior angle formed by
junction of radius and humerus; ventral surface of wing membrane
furred from just behind anterior border to a line extending from
knee joint to basal third of third metacarpal. The fur constituting
this strip is thicker and longer at the base of the fifth metacarpal
than elsewhere. Whole dorsal surface of interfemoral membrane
furred; but its ventral surface furred only at extreme base. Dorsal
surface of toes furred.
Color.—Hairs black at bases, except on face, chin, and membranes.
In general, hairs on face and chin are yellowish, tipped with rufous.
Hairs on back black for 1.5 millimeters from base; sueceeding 7 milli-
meters pale yellowish; distal to this a rufous band about 1 millimeter
wide, followed by an ashy tip of 0.5 millimeter. Ashy tips often
lacking on posterior portion of back, as also on dorsal surface of inter-
femoral membrane. On the latter area the black at bases of the hairs
may also be wanting.
IT am informed by Messrs. Miller and Hollister of the United States
National Museum that in the eastern red bat the males are, on the
average, brighter colored than the females, it being possible in most
specimens to determine the sex by color alone. I find that this does
not hold true with our western subspecies. McAtee (1907, p. 8) men-
tions the resemblance of an individual of the eastern red bat to a
withered leaf caught among the twigs of a tree. This suggests a
protective value for the reddish brown color of our species.
nia 325
f the Bats of Califor
: A Synopsis o
Grinnell
1918]
MEASUREMENTS IN MILLIMETERS OF ELEVEN
Mus.
no.
3277
4048
4049
4674
5146
6958
18794
16598
16670
21438
24327
Sex
+0 +0 +0 +0 Og Og A OY AA OG OW
Locality
Berkeley, Alameda Co.
Westley, Stanislaus Co.
Westley, Stanislaus Co.
Napa, Napa Co.
Pasadena, Los Angeles Co.
Glendora, Los Angeles Co.
Escondido, San Diego Co.
Live Oak, Sutter Co.
Cuyama Valley, San Luis Obispo
Fresno, Fresno Co.
Fresno, Fresno Co.
Specimens or NYCTERIS BOREALIS TELIOTIS (H. ALLEN), FROM CALIFORNIA
Co.
106.0
99.0
105.0
109.0
111.0
110.0
vertebrae
Tail
eranium
ygomatic
breadth
a)
wo OZ
cS)
re)
“ Breadth of
NNAN
om NO © bprain-case
a
fo)
7.5
7.5
7.7
7.5
8.0
= Interorbital
constriction
hes ee obra
SRO
n
4.5
4.3
326 Umiversity of California Publications in Zoology [Vou.17
Skull—As described for the genus. May always be distinguished
from that of Nycteris cinerea, by its decidedly smaller size (pl. 23,
fig. 51, and pl. 24, fig. 59). The average skull length of the examples
of cinerea in the Museum of Vertebrate Zoology is 17.5 millimeters,
while that of the series of teliotis is but 12.5 millimeters.
Measurements.—Average and extreme measurements in milli-
meters of a series of 11 specimens in the Museum of Vertebrate
Zoology are as follows: Total length, 104.5 (extremes, 98.0-111.0) ;
tail vertebrae, 50.5 (42.0-60.0); tibia, 18.5 (17.0-19.7); foot, 8.5
(6.0-11.0) ; forearm, 38.7 (34.0-41.3) ; greatest length of skull, 12.5
(12.0-13.1).
Synonymy and History—This bat was described by H. Allen
(1891, pp. 5-7) under the name Atalapha teliotis. The description was
based on a poor specimen without data, but presumably taken in
southern California. This type is now number 84555, in the United
States National Museum (Lyon and Osgood, 1909, p. 277).
Distribution—The general range of Nycteris borealis teliotis is
given by Miller (1912, p. 64) as follows: ‘‘From the head of Sacra-
mento Valley, California, south to Comondu, Lower California.’’
J. Grinnell (19130, p. 280) gives the California range in winter and
spring as Sacramento and San Joaquin valleys, from Sutter County
southwards, and throughout the San Diegan district. (See map,
text-fig. R.) A study of the dates of capture of California red bats
leads the present writer to infer that the sexes separate during the
summer months, the females remaining in the Lower Sonoran zone,
while the males migrate into the Upper Sonoran and Transition zones.
Among certain species of birds the males withdraw from the
breeding grounds in early summer and forage elsewhere, often at
higher elevations. Their departure relieves congestion in the nesting
area and leaves a greater food-supply for the females and young.
A study of the summer distribution of red bats in California leads
one to believe that a similar habit exists among these vertebrates
also. Of the red bats listed, with dates of capture, in the accompany-
ing table, the nineteen adult females taken between April 15 and
August 1 are all recorded from the Lower Sonoran zone; whereas
of the four adult males taken during the same time of the year
three are from the Transition zone and the fourth from the Upper
Sonoran. All winter records of red bats, of both sexes, are from the
two Sonoran zones. :
Specimens Examined.—The writer has examined 56 examples of
this bat from California. Since the seasonal movements of the Cali-
fornia red bat have not been fully worked out, it seems pertinent to
1918] Grinnell: A Synopsis of the Bats of California 327
give here the date as well as locality of capture of each specimen.
The entire list appears on an accompanying table.
Natural History —The western red bat is a solitary, tree-dwelling
species. It spends its days hanging in foliage. Stephens (1906,
nN
OAPAN i Pk
pri Eek)
@ NYCTERIS BOREALIS TELIOTIS
& NYCTERIS CINEREA
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNLA,
Fig. R. Map showing stations of occurrence in California of Nycteris borealis
teliotis and Nycteris cinerea, as established by specimens examined by the author.
p. 271) states that all that he has seen were found in spring or summer
hanging in fruit trees in orchards.
One of the western red bats in the Museum of Vertebrate Zoology
(no. 18794) was found hanging among the evergreen leaves of an
orange tree near Escondido, San Diego County, October 8, 1912.
328
University of California Publications in Zoology
[Vou. 17
TaBLeE SHow1ne Locavities (ARRANGED FROM NorTH TO SoUTH) AND DATES OF
CAPTURE OF NYCTERIS BOREALIS TELIOTIS (H. ALLEN) IN CALIFORNIA
No
Sex
65375 Biol. Surv. dad.
23897
16599
16600
16598
4674
6974 Amer. Mus.
11675 Phila. Acad.
24176
24177
24178
24179
3277
132344 Biol. Surv.
23898
U. C. Dept. Zool.
105250 Nat. Mus.
Stanford Univ.
Stanford Univ.
Stanford Univ.
Stanford Univ.
Stanford Univ.
Stanford Uniy.
Stanford Uniy.
4048
4049
21438
21514
21515
21516
21902
21903
21904
21905
21906
21907
21908
21909
21910
21911
21912
21913
21914
g
9 ad.
9 ad.
ad.
od ad.
dad.
@ ad.
9 juv.
J ad.
° ad.
Jo ad.
dad.
dad.
dad.
dad.
dad.
@ ad.
© ad.
© ad.
° ad.
° ad.
© ad.
° ad.
9 ad.
? ad.
° ad.
3 juv.
of juv.
3 juv.
9 ad.
3 juv.
3 juv.
3 juv.
Locality
Tehama Co.: Tehama
Mendocino Co.: near
Laytonville
Yuba Co.: Hammonton
Yuba Co.: Hammonton
Sutter Co.: Live Oak
Napa Co.: Napa
Marin Co.: Nicasio
San Joaquin Co,: Linden
San Joaquin Co.: Stockton
San Joaquin Co.: Stockton
- San Joaquin Co.: Stockton
San Joaquin Co.: Stockton
Alameda Co.: Berkeley
Alameda Co.: Haywards
Alameda Co.: Berkeley
Alameda Co.: Berkeley
San Mateo Co.: Colma
Santa Clara Co.: Palo Alto
Santa Clara Co.: Palo Alto
Santa Clara Co.: Stanford
University
Santa Clara Co.:
University
Santa Clara Co.:
University
Stanford
Stanford
Santa Clara Co.: Palo Alto
Santa Clara Co.: San Jose
Stanislaus Co.: Westley
Stanislaus Co.: Westley
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Fresno Co.: Fresno
Date
May 14, 1894
Apr. 2,1916
. 15, 1912
3, 1912
15, 1908
26, 1891
2, 1898
12, 1915
12, 1915
12, 1915
12, 1915
22, 1903
. 28, 1904
8, 1916
Oct.
Aug.
June
July
July
July
July
17, 1898
. 27,1900
7, 1893
12, 1910
1893
Jan.
Sept.,
Sept., 1893
May 10, 1900
Oct.
Feb.,
Feb.
Feb.
Apr.
May
May
May
May
May
May
May
May
June
June
June
June
June
June
June
June
3, 1893
1897
12, 1909
23, 1909
3, 1915
21, 1915
21, 1915
21,1915
28, 1915
28, 1915
28, 1915
28, 1915
28, 1915
23, 1915
23, 1915
23, 1915
23, 1915
23, 1915
23, 1915
23, 1915
23, 1915
. 15,1912"
Collector
C. P. Streator
F. C. Clarke
Mrs. F. H. Holden
Mrs. F. H. Holden
Mrs. F. H. Holden
Mrs. F. H. Holden
A. S. Bunnell
W. O. Emerson
Mrs. J. L. Schlis-
inger
Miss Reed
. Hornung
. G. Buxton
. O. Snyder
. M. Stowell
SUAS
J. M. Stowell
O. Jenkins
W. W. Price
A. W. Greely
. Brookey
. Brookey
Borell
Borell
Borell
Borell
and R.
and R.
and R.
and R.
and R.
Borell
Borell
Borell
Borell
Borell
Borell
. Borell
. Borell
Borell
Borell
Borell
Borell
Borell
HHH EPR
Grinnell: A Synopsis of the Bats of California
1918] 329
No. Sex Locality Date Collector
31177 Biol. Surv. gad. Tulare Co.: Three Rivers Sept. 16,1891 A. K. Fisher
16670 Q ad. San Luis Obispo Co.: Apr. 22,1912 J. Grinnell
Cuyama Valley
54602 Biol. Surv. Santa Barbara Co.: May 20,1893 H.K. Chamberlain
Santa Barbara
30729 Biol. Surv. @Qad. Kern Co.: Bakersfield July 17,1891 V. Bailey
6958 gad. Los Angeles Co.: Glendora Dec. 10,1903 W. 8. Wood
187714 Nat. Mus. dad. Los Angeles Co.: Alhambra Dec. 28,1880 E. C. Thurber
Coll. F. Stephens @ ad. San Bernardino Co.: May 15,1903 F. Stephens
Warren’s Ranch
Stanford Univ. gad. Riverside Co.: San Jacinto Noy. 15,1893 E. Hyatt
18794 dad. San Diego Co.: Escondido Oct. 8,1912 J. Dixon
2515 Coll. F. dad. San Diego Co.: Witch May 25,1895 F. Stephens
Stephens Creek
62870 Nat. Mus. San Diego Co.: Witch June 19,1895 A. W. Anthony
Creek
716 Coll. F. Gad. San Diego Co.: Santa Aug. 15,1896 EF. Stephens
Stephens Ysabel
60533 Nat.Mus. dad. San Diego Co.: Santa June 15,1892 F. Stephens
Ysabel
Another (no. 6958) was found hanging in an orange tree at Glendora,
Los Angeles County, December 10, 1903. The female red bat (no.
21488) secured by Adrey Borell at Fresno, April 3, was found hang-
ing in a mulberry tree in company with ‘‘many’’ more of its kind and
one hoary bat, which was also captured.
In some portions of its general range the red bat is migratory,
while in others it is said to hibernate in vast numbers in caves. The
latter report needs to be verified. This subject is treated more fully
under the heading Migration, on page 282.
In most bats the number of mammae is two, but in the bats of the
genus Nycteris the number is four. This fact might lead to the
inference that in this genus there is a greater number of young than
in other genera. Observations made by Lyon (1903, pp. 425-426)
and Ward (1905, p. 20) establish the fact that as many as four young
oceur at a single birth. Lyon records the capture at Washington,
D. C., of a female eastern red bat weighing 11 grams. Clinging to
her nipples were four young whose combined weights were 12.7
grams. Combining his own information with that given by Lyon,
Ward gives the following summary of observations upon the number
of young clinging to, or embryos found in, adult females of the
genus Nycteris: two with one, two with two, three with three, two
with four.
330 University of California Publications in Zoology (|Vou.17
Among the female western red bats in the Museum of Vertebrate
Zoology are five which contained embryos, and three captured with
young. Of the former, four contained three embryos each and the
fifth, two. Each of the three mothers had three young clinging to
-her. Two of the mothers were found in adjoining peach trees at
Fresno, June 23, 1915. Of these two, no. 21907, when discovered,
had three small naked young clinging to her, all males; the young
clinging to no. 21911 were also all males, but were twice the size of
the individuals in the other family. The third mother, no. 24179,
was found at Stockton, on July 12, 1915. The bat was clinging to
the trunk of a tree one and one-half feet above the ground, with
her young attached. Her weight was 12.4 grams and the weights
of her three half-grown young were: two males, 6.5 each; one female,
6.4; total, 19.4 grams. (See pl. 18, fig. 18.)
Nycteris cinerea (Peale and Beauvois)
Hoary Bat
Vespertilio cinereus Peale and Beauvois (1796, p. 15). Original deserip-
tion; type locality, Pennsylvania.
‘Lasiurus cinereus, H. Allen (1864, pp. 21-24, figs. 18-20). Description;
specimens recorded from Petaluma and Monterey.
Lasiurus cinereus, Cooper (1868, p. 5). Distribution.
L{asiurus]. cinereus, Cooper (in Cronise, 1868, p. 442). Occurrence in
California.
Atalapha cinerea, Dobson (1878, pp. 272-274). Description; specimen
listed from Monterey.
Atalapha cinerea, Bryant (1891a, p. 358). Nominal.
Atalapha cinerea, H. Allen (1894, pp. 155-162, pls. 29-31). Description.
Lasiurus cinereus, Miller (1897b, pp. 112-115, figs. 31-82, pl. 3, fig. 4).
Description; distribution; lists several California localities.
Lasiurus cinereus, Elliot (1901, p. 413). Diagnosis; general distribution.
Lasiurus cinereus, Stone (1904b, p. 587). Record of occurrence at Berke-
ley.
Lasiurus cinereus, Elliot (1904b, p. 595, fig. 88). Diagnosis; general dis-
tribution. ;
Lasiwrus cinereus, Stephens (1906, p. 272). Deseription; distribution ;
habits.
Lasiurus cinereus, Elliot (1907, pp. 513-514). Record of specimen taken
at Nicasio.
Lasiurus cinereus, Seton (1909, p. 1193). Map showing record stations
and hypothetical range.
Nycteris cinerea, Miller (1912, p. 64). General range.
Nycteris cinerea, J. Grinnell (1913), p. 280). Range in California.
Diagnosis—Size large (forearm 49 to 56.6 millimeters long) :
whole dorsal surface of interfemoral membrane furred; color, yellow-
ish brown, conspicuously tipped with silvery white.
1918 | Grinnell: A Synopsis of the Bats of California 331
Description: Head.—Muzzle broad; greatest width of rostrum
about one-half entire length of head (pl. 16, fig. 8). Nostrils direeted
obliquely outward and slightly downward, and their rims widely
diverging, tumid; space between them concave and, in dried skins,
their openings more than 3 millimeters apart. Ear short, about 18
millimeters in height from meatus, and broad, only 1 millimeter less
wide than high; external basal lobe about size of tragus and not
notched on its anterior border; tragus somewhat triangular in outline
and about half height of ear from crown.
LTimbs and Membranes—Wings unusually lone (extent almost
three times total length of body) and narrow; third metacarpal ex-
ceeding fifth by at least length of thumb (pl. 16, fig. 7). Wings and
interfemoral membranes attached at bases of toes. Tip of tail in-
eluded in interfemoral membrane; tail unusually long, length of
caudal vertebrae exceeding that of forearm by from 4 to 11 milli-
meters.
Pelage.—F ur similar to that of red bat, everywhere full and soft,
and longest on dorsal surface between shoulders, where from 10 to
12 millimeters in length. Outer side of ear furred on lower half; a
strip of hair near anterior edge of ventral surface of ear; a squarish
patch of hairs on ventral surface of ear in front of apex; basal lobe
furred on outside. Tragus scantily haired on outer surface. Dorsal
surface of wings furred as follows: a line of fur, continuous with
that of body and of dorsal surface of interfemoral membrane, extends
from ankle along surface of wing to anterior edge of antebrachial
membrane opposite middle of humerus (pl. 17, fig. 10); along each
side of radius is a scanty growth of short hairs, averaging less than
1 millimeter in length, except for oblong patch of longer and denser
fur on proximal fifth of forearm on edge nearest body. Another
pateh of fur les at the proximal end of the fifth metacarpal; from
this a scanty line of hairs extends along sides of metacarpal for one-
third its length. A third patch of fur lies at base of thumb. On
ventral surface of wing a line of fur continuous with that of body
extends from knee joint to distal end of radius, where it fills the
angles between the finger joints and extends outwardly along inner
edge of third metacarpal for two-thirds its length. Ventral surface
of antebrachial membrane furred except at its anterior edge. Dorsal
surface of interfemoral membrane entirely furred, as also the dorsal
surfaces of the toes.
Color.—Dorsal surface of rostrum, all hairs on ears, a band about
8 millimeters wide extending across throat and uniting ears, and
fur on under surface of wings, near naples yellow; fur on wings
nearest body, with brownish base. Sides of muzzle, rims of ears,
and chin, blackish brown. Hairs on whole back and dorsal surface
of tail seal brown for about 2 millimeters at bases, then light buff
for another 2 or 3 millimeters, followed by seal brown for about 1.5
millimeters, and terminated by a white tipping on the distal end of
each. Bands of buff are missing from hairs on dorsal surface of tail.
On ventral surface of body a ruff of hairs, similar to those on
back, encircles throat just posterior to the band of naples yellow.
Posterior to this the hairs have bases of light seal brown, and tips
of light buff. On the dorsal surface of the wing the three small
patches of fur (2 millimeters in diameter) are pale buff in color.
332 University of California Publications in Zoology {|Vou.17
Wing membranes nearest chaetura black, except area on dorsal
surface directly above furred portion of ventral surface, which is
light cinnamon brown; specklings of this color extend out onto the
darker portions of membranes.
According to Miller (1897), p. 113) the color variation, although
considerable, is never enough to obscure the characters of the species
and appears to be wholly independent of locality, certain skins from
such widely separated regions as Minnesota and southern California
being practically indistinguishable.
Skull—sSimilar to that of Nycteris borealis teliotis but larger
(pl. 28, fig. 49, and pl. 24, fig. 57). In N. b. teliotis the skull length
varies from 12 to 13.1 millimeters, while in N. cinerea it is from 16.9
to 18.5 millimeters.
Measurements—Average and extreme measurements in milli-
meters of a series of 14 skins in the Museum of Vertebrate Zoology
are as follows: Total length, 135.1 (extremes 128.0-146.0) ; tail verte-
brae, 57.7 (51.0—-62.0) ; tibia, 20.4 (19.0-22.5) ; foot, 11.1 (10.0-12.0) ;
forearm, 51.8 (49.0-56.6) ; greatest leneth of skull 17.5 (16.9-18.5).
Synonymy and History—The hoary bat was first deseribed by
Peale and Beauvois (1796, p. 15) from a specimen secured in Penn-
sylvania, probably near Philadelphia.
Distribution—The general range of this bat is given by Miller
(1912, p. 64) as extending throughout ‘‘boreal North America from
the Atlantic to the Pacific, breeding within the Boreal Zone, but in
autumn and winter migrating at least to the southern border of the
United States.’’ The range in California in winter and spring
comprises the valleys of west-central and southern California, south
through the San Diegan district; in summer probably the Transition
and Boreal zones. (See map, text-fig. R.)
The accompanying table shows very few strictly summer records
of the hoary bat in California. It is possible that many of the indi-
viduals wintering here come from breeding areas north of this state.
A discussion of the migration of this bat is given in the present paper
under the heading Migration, on page 232.
Specimens Examined.—The writer has examined 55 examples of
the hoary bat from California. Since the seasonal movements of this
species are not fully known the writer has included in the accompany-
ing table the dates of capture as well as the localities from which
specimens have been secured.
Natural History.—Several hoary bats have been found hanging in
the thick foliage of orange trees in southern California in the winter.
On April 17, 1904, H. 8. Swarth discovered one clinging to a branch
of an oak tree, looking like some huge gall. Stephens found the
species in the redwoods of Mendocino County in May. He records
3
€
e
€
3
e
3
ub
f the Bats of Californ
A Synopsis o
Grinnell
1918]
MEASUREMENTS IN MILLIMETERS OF FOURTEEN SPECIMENS OF NYCTERIS CINEREA (PEALE AND BEAUVOIS), FROM CALIFORNIA
aannaaara#araanaandan Interorbital
2 “ 2 38
no. Sex Locality (=fs' Baia ry = £ ein Se a7
5147 3 Nordhoff, Ventura Co. 130.0 57.0 19.6 11.0 51.0 17.4 12.6 10.2
6944 fof Glendora, Los Angeles Co. 135.0 59.0 19.0 10.0 52.2 17.2 12.0 10.0
9467 Jb Los Angeles, Los Angeles Co. 140.0 HYsHO! Sees 11.0 54.0 18.5 13.2 11.0
14650 3 Raymond, Madera Co. 128.0 51.0 19.0 11.0 50.0 17.3 12.4 10.0
14651 3 Raymond, Madera Co. 130.0 55.0 21.6 11.0 51.0 17.4 12.3 10.1
14652 fof Raymond, Madera Co. 137.0 57.0 22.5 11.0 5010 ee 12.8 10.3
20778 Jb Berkeley, Alameda Co. 140.0 G220ge 12. 51.3 17.4 12.3 10.1
21311 fed Berkeley, Alameda Co. 129.0 58.0 11.5 BURBI) E arae. 9 peetese uae | ees
20815 Jb 1 mi. west Guerneville, Sonoma Co. 131.0 610 geeess 12.0 51.5 17.2 12.6 10.0
3581 2 Monterey, Monterey Co. z 145.0 59.0 21.0 12.0 56.6 aRBs}s eee 10.0
21439 g Fresno, Fresno Co. 132.0 59.0 20.7 11.0 49.0 17.9 13.0 10.3
6945 2 San Fernando Valley, Los Angeles Co. 131.0 58.0 20.0 11.0 50.4 16.9 12.1 10.1
5148 2 Pasadena, Los Angeles Co. 138.0 58.0 20.2 10.0 51.5 17.8 12.7 10.3
5083 2 Dinuba, Tulare Co. 146.0 Gil Oe 12.0 55.5 17.8 12.3 10.3
constriction
wHwt tH & fb hw Ob PP
334
University of California Publications in Zoology
TaBLE SHOWING LOCALITIES (ARRANGED FROM NortTH TO SOUTH) AND DATES
oF CAPTURE OF NYCTERIS CINEREA (PEALE AND BEAUVOIS) IN CALIFORNIA
No.
16651 Biol. Surv.
98095 Biol. Surv.
711 Stanford Univ.
1101 Amer. Mus.
1099 Amer. Mus.
187824 Nation. Mus.
187825 Nation. Mus.
22301 Nation. Mus.
59988 Nation Mus.
187822 Nation. Mus.
187823 Nation. Mus.
20815
16516 Biol. Surv.
20778
11673 Phila. Acad.
U. C. Dept. Zool.
U. C. Dept. Zool.
U. C. Dept. Zool.
21312
U. C. Dept. Zool.
119948 Biol. Surv.
105259 Nation. Mus.
331 Stanford Univ.
9406 Mus. Comp.
Zool.
1438 Stanford Univ.
1437 Stanford Univ.
Stanford Univ.
Stanford Univ.
Stanford Univ.
199608 Nation. Mus.
Stanford Univ.
Calif. Acad. Sci.
23899
23039
14650
14651
14652
21439
17058 Biol. Sury.
Sex
Ch fowl H Cyr, Chie
POON ON TON OS
+0 +0 O, O, Q, +0:
Locality
Humboldt Ca.:
Haydenhill
Lassen Co.:
Mendocino Co.:
Nicasio
Nicasio
Nicasio
: Nicasio
: Nicasio
: Nicasio
: Nicasio
: Nicasio
1 mi. W.
Co.:
Co.:
Co.:
Marin
Marin
Marin
Marin
Marin
Marin
Marin
Marin
Sonoma Co.:
Guerneville
Sonoma Co.:
Alameda
Alameda
Alameda
Alameda
Alameda
Alameda
Co.:
Co.:
Co.:
Alameda Co.:
Alameda Co.:
San Francisco
Francisco
2)
Santa Clara Co.:
University
Santa Clara Co.:
Santa Clara Co.:
University
Santa Clara Co.:
University
Santa Clara Co.:
Santa Clara Co.:
Santa Clara Co.:
Santa Clara Co.:
Santa Clara Co.:
View
Santa Clara Co.:
View
Eureka
Cahto
Cloverdale
: Berkeley
: Berkeley
: Berkeley
Berkeley
Berkeley
Berkeley
Berkeley
Haywards
Co.: San
Stanford
Palo Alto
Stanford
Stanford
Menlo Park
Santa Clara
Menlo Park
Gilroy
Mountain
Mountain
Merced Co.: Snelling
Mariposa Co.:
Madera Co.:
Madera Co.:
Madera Co.:
Merced Lake
Raymond
Raymond
Raymond
Fresno Co.: Fresno
Santa Cruz Co.
: Santa Cruz
May
July
May
Oct.
Oct.
Oct.
Oct.
Sept.
Sept.
Sept.
Oct.
July
Apr.
Oct.
Date
25, 1889
16, 1899
24, 1889
17, 1887
5, 1887
19, 1888
25, 1888
4, 1891
15, 1891
14, 1888
13, 1888
11,1913
18, 1889
6, 1914
7, 1898
6, 1905
1, 1912
5, 1905
10, 1915
7, 1900
6, 1902
1, 1909
. 27, 1909
. 11, 1898
27, 1894
. 28, 1894
23, 1893
1, 1893
31, 1894
6, 22, 1915
3, 1896
29, 1905
. 15, 1916
. 21, 1915
5 ip alenlal
17,1911
. 21,1911
3,(1915
5, 1910
[Vou. 17
|
|
Collector
T.S. Palmer
. Bailey |
C. MeGregor
. Allen
. Allen
. Allen
. Allen
. Allen |
. Allen |
Allen
. Allen
and H. W.
Grinnell
T.S. Palmer
Sam Brodie
A.S. Bunnell
uogagaaaaps
PP PP PP PP
W. C. New-
berry
W. O. Emerson
J. Hornung
J. Dixon
W. W. Price
J. VanDen-
burgh
— Magee
J. M. Hyde
F. G. Krauss
H. Doud
E. M. Ehrhorn
H. O. Jenkins
J. Grinnell
©. A. Gallo
J. Grinnell
J. Grinnell
J. Grinnell
Adrey Borell
C. P. Streator
1918 | Grinnell: A Synopsis of the Bats of California 335
No. Sex Locality Date Collector
3581 2 Monterey Co.: Monterey Noy. 10,1907 J. Rowley
5033 io) Tulare Co.: Dinuba Apr. 1, 1909 A. L. Dickey
27977 Biol. Surv. 2 Inyo Co.: Panamint Mts. May 17,1891 E.W. Nelson
28946 Biol. Surv. do Inyo Co.: Panamint Mts. Apr. 21,1891 E. W. Nelson
5147 3d Ventura Co.: Nordhoff Jan. 18,1905 J. Grinnell
29845 Biol. Surv. 3 Kern Co.: 25 mi. above July 6,1891 <A.K, Fisher
Kernville
5148 Q Los Angeles Co.: Pasadena Feb. 5,1906 J. Grinnell
3423 = Los Angeles Co.: Los May ...., 1890 E. C, Thurber
Angeles
6944 3 Los Angeles Co.: Glendora Dee. 29,1906 J. Grinnell
6945 2 Los Angeles Co.:; San Fer- May 7,1904 J. Grinnell
nando Valley
9467 3 Los Angeles Co.: Los Apr. 17,1904 H.S.Swarth
Angeles
8248 Phila. Acad. 5: San Bernardino Co.: San Apr. 24,1893 R.B. Herron
Bernardino Valley
125801 Biol. Surv. ef San Bernardino Co.: War- May 15,1902 F. Stephens
ren’s Ranch
Coll, F. Stephens go Riverside Co.: Riverside Feb. 21,1889 F.0O. Johnson
Coll. F. Stephens 2 San Diego Co.: San Diego Dee. 20,1903 FE. Stephens
53810 Biol. Sury. = San Diego Co.: Santa Ysabel May 10,1893 H.W. Hyatt
the flight as swift, with frequent abrupt turns, and states that these
bats do not appear until the light becomes very dim.
A female hoary bat secured at Merced Lake, Mariposa County, was
shot at 7:12 p.m. on the cloudy evening of August 21, 1915, as it
flew among the lodgepole pines at a height of about thirty feet above
the ground.
Merriam (1884, pp. 176-181) gives an interesting account of this
bat as occurring in the Adirondack region of New York state. There
the hoary bat was the latest of the observed species to appear in the
evening. Merriam states that this bat may be recognized in the dark
by its great size, long and pointed wings, and the swiftness and
irregularity of its flight.
Seton (1909, p. 1169) states that the hoary bat is the only known
species which is never gregarious. As occurring in California it is
certainly a notably solitary species. The only evidence to the con-
trary is afforded by the female hoary bat secured at Fresno, April 3,
1915, which was found hanging in a mulberry tree is company with
“‘many’’ red bats.
Seton (1909, p. 1196) states that four is the usual number of
young at birth. Banta (in McAtee, 1907, p. 8) states that there is
in the museum at the University of Indiana a female hoary bat which
when captured was found to have two young clinging to it.
336 University of California Publications in Zoology |Vou.17
Genus Euderma H. Allen
This genus, as at present known, consists of a single species, found
only in the southwestern United States.
Characters.—Kars very large, about three-fourths as long as fore-
arm, joined across forehead by band of membrane; nostrils simple,
facial portion of skull narrow and pointed, the brain-case quadrate,
flattened above, but rising abruptly at frontal border; zygomata
abruptly expanded at middle; auditory bullae very large, and much
elongated, their greatest diameter about equal to length of tooth-row
exclusive of incisors.
2a =i 2-2 3-3
3-37 “1-1 P™ 9-9 ™ 3-3
cisors alike in form, slender, with well developed cingula, and simple
styliform crowns, the inner somewhat larger than the outer, which
is In contact, or nearly so, with canine. Lower incisors trifid, middle
lobe largest, especially in second and third. Upper canine small
and weak, barely more than twice height of inner incisor, which it
rather closely resembles in form. Lower canine relatively smaller
than in any other known Vespertilionine bat, its crown searcely higher
than that of fourth lower premolar. Anterior upper premolar
minute, scarcely more than one-third as large as outer incisor and
rising barely to cingulum of canine; in the tooth-row it stands at the
middle of the narrow space between the canine and large premolar,
and in form resembles the incisors, but its crown is relatively lower
and thicker; large premolar of usual form, but very deeply coneave
on anterior border. Lower premolars of ordinary Vespertilionine
type; molars normal. [As no skull of Euderma is at hand, the above
description has been adapted from Miller (1907, pp. 225-227). |
Comparisons.—In length of ears and form of skull and teeth this
genus is said to resemble Plecotus and Corynorhinus, but its simple
nostrils are of the Myotis type.
Dental formula: i
= 34. Upper in-
Euderma maculatum (J. A. Allen)
Spotted Bat
Histiotus maculatus J. A. Allen (1891a, pp. 195-198). Original deserip-
tion; type locality near Piru, Ventura County, California.
Histiotus maculatus, Bryant (1891b, p. 113). Nominal.
Histiotus maculatus, H. Allen (1892, pp. 467-470). As type of the new
genus Huderma.
Buderma maculatum, Bryant (1892, p. 220). Nominal.
Euderma maculata, H. Allen (1894, pp. 61-64). Description.
Euderma maculatum, Miller (1897b, pp. 46-49, pl. 3, fig. 3). Description.
Euderma maculata, Elliot (1901, p. 398). Description; distribution.
Euderma maculatum, Miller and Rehn (1901, p. 265). Type locality.
Euderma maculatum, Elliot (1904b, p. 283). Reference to type locality.
Euderma maculatum, Elliot (1905, p. 490). Distribution.
Euderma maculatum, Stephens (1906, p. 264). Description; distribution.
1918] Grinnell: A Synopsis of the Bats of California 307
Euderma maculatum, Miller (1907, pp. 226-267, 1 fig.). Description of
genus.
Euderma maculatum, J. Grinnell (1910, pp. 317-320, pl. 30). Second record
from California.
Euderma maculatum, Miller (1912, p. 67). Nominal.
ELuderma maculatum, J. Grinnell (1913b, p. 281). Range in California.
Diagnosis —Same as for the genus.
Description: Head—Kars (pl. 16, fig. 9) very large, and in the
dried skin at hand widely diverging at tips, fully three-fourths as
long as forearm, and joined across forehead by a low band of mem-
brane; ear convex on inner border, broadly rounded at tip, slightly
concave on outer border just below tip, and convex on basal half of
outer border, which continues as a low fold to a point below angle of
mouth. Tragus well developed, 19 millimeters in length (in dried
skin), this being more than one-third length of ear; posterior border
convex above small basal lobe, tip broadly rounded, and anterior
border straight on upper half, but slightly convex below. As sug-
gested by Miller (1897b, p. 47) the tragus resembles in form the blade
of a table knife. Nostrils prominent and situated at end of a narrow,
low, naked dise, divided by a slight groove, and narrowing posteriorly
to a point.
Limbs and Membranes.—Forearm slightly bowed; thumb with a
small basal pad. Wing short and stout, length of third metacarpal
scarcely exceeding that of fifth. Wing and tail membranes wholly
devoid of hair. About half of last caudal vertebra exserted. Feet
moderately large, a little less than half as long as tibia. Toes very
seantily haired.
Pelage—F ur soft, silky, and about 8 millimeters in length, save
for a small woolly patch at posterior base of each ear.
Color.—The small woolly patches of fur at bases of ears are whitish
throughout. Elsewhere on body all hairs are blackish either on the
proximal third or for their entire length. On the dorsal surface
of the animal are three equal-sized white patches of fur about 9 by 17
millimeters in extent, one on each shoulder, and one on the rump
(pl. 16, fig. 9). As suggested by J. Grinnell (1910, p. 318) these
remind the observer of the ‘‘death’s-head’’ pattern of certain moths.
The hair in these patches is, as elsewhere, blackish at base. On the
ventral surface, the distal third of the hair is everywhere white, save
for a narrow collar of blackish hairs continuous with blackish fur
of back. In a dried skin which has been preserved for five years the
ear membranes are yellowish, other membranes pale brown.
Skull—As given for the genus.
Measurements——The accompanying table of measurements has
been compiled from published descriptions of specimens.
Synonymy and History.—This bat was first described by J. A.
Allen (1891a, pp. 195-198) from a single specimen found ‘‘ caught
on a fence at Piru, in the western part of Ventura County, Califor-
nia.’’ This specimen was secured by a Los Angeles taxidermist,
Thomas Shooter, and through E. C. Thurber transmitted to the
338 University of California Publications in Zoology {Vou.17
MEASUREMENTS IN MILLIMETERS OF THREE EXAMPLES OF EUDERMA MACULATUM
(J. A. ALLEN)
= 2
Fe B E 33
=h = s s Es
Mus £8 ms ra S Fa ae
no. Sex Date i ape = <= £ Ki
1196* ? Oct.1; 1907s" ese) Bee 18.0 45:0) ‘a2 “eS
1225457 diad. Sept., 1903 107.0 47.0 196 9.8 496 188 104
3920
By a March, 1890 110.0 50.0 21.0 98 500: 19.0 10.9
1From Mecca, Riverside County, California.
2U. S. National Museum: from Mesilla Park, New Mexico.
3 Amer. Mus. Natural History: from Piru, Ventura County, California.
American Museum of Natural History. Dr. C. Hart Merriam (in
Miller, 18976, p. 49) gives the added information that the type was
in reality taken at the mouth of Castae Creek, in the Santa Clara
Valley; this is eight miles from Piru, in the same valley, but across
the line in Los Angeles County.
Because of its general resemblance, as regards size, and the form
and size of the ears, to Histiotus of South America, Dr. J. A. Allen
placed the bat in that genus. H. Allen (1892, pp. 467-470) placed
the bat in the new genus Huderma, separating it from Histiotus upon
the presence of a minute first upper premolar, absent in the latter
genus.
Distribution.—The spotted bat is known from only four localities,
in each of which but a single individual has been secured. The
localities and dates are as follows: near Piru, Ventura County, Cali-
fornia (see above), the type, March, 1890 (J. A. Allen, loc. cit.) ;
Mecea, Imperial County, California, an individual found dead in
the overflow from a railway watering tank, October 1, 1907 (J. Grin-
nell, 1910, p. 317) ; Mesilla Park, New Mexico, an individual found
dead in the biological laboratory of the New Mexico College of Agri-
cultural and Mechanical Arts, in September, 1903 (Miller, 1903,
p- 165) ; Yuma, Arizona, one reported by Herbert Brown (Stephens,
1906, p. 264). (See map, text-fig. O.)
Dr. C. Hart Merriam (in Miller, 1897b, p. 49) states that while
in Vegas Valley, Nevada, he was told of the occurrence there, during
the summer, of a very large bat ‘‘with ears like a jackass and a white
stripe on each shoulder.”’
Specimens Examined.—The writer has seen but one example of
Euderma maculatum from California: Mecca, Riverside County, 1.
Natural History Nothing is known of the habits of this species.
1918] Grinnell: A Synopsis of the Bats of California 339
Genus Corynorhinus H. Allen
This genus is found only in the warmer portions of North America,
from southern British Columbia to southern Mexico. Three species
are now recognized, one of which is divisible into four races.
Diagnosis—KEars very long, about three-fourths length of fore-
arm; tragus slender, pointed at tip and wholly free from external
basal lobe of ear.
Description: Head—Muazzle slender; greatest width of rostrum
less than one-third total length of skull. This slenderness, however,
is somewhat concealed by club-shaped glandular masses which rise
on each side of snout, between nostril and eye, to a height of 2.5
millimeters (in an alcoholic specimen) above sur-
face of rostrum, the two of which converge nearly
to meet in the median line. Nostrils irregularly
quadrate and placed on sides of muzzle, opening
outwards and slightly upwards; borders of nostrils
slightly thickened, giving them a rimmed appear-
ance. Ears very large (text-fig. S), being about
three-fourths length of forearm, and joined across
forehead by a narrow band of membrane. Tragus
straight and slender, about two-fifths height of ear.
Anterior border of auricle strongly convex and
turned abruptly backward so as to rest upon dorsal Fig. &. Side
surface of ear almost to its bluntly rounded tip. view of head of
Posterior half of ear marked by a series of trans- Corynorhinus — raf-
inesquit pallescens
verse velnings, most prominent near center of ear
and fading away towards its posterior border.
(See pl. 15.)
(drawn from speci-
men no. 9368), X
1.00, showing long
ear, long acutely
pointed tragus,
and glandular
swelling on side of
muzzle.
Skull and Teeth—Skull slender, with com-
pressed zygomata ; brain-case elongate and rounded,
in profile shghtly depressed on top and convex
anteriorly; rostrum coneave in profile, sloping
abruptly downward from its junction with brain-
ease. Auditory bullae large and rounded; their greatest diameter
equal to about three times distance between them. (See pl. 23, fig. 50;
pl. 24, fig. 58.)
Mental cf Te Bare lke 2-2 3-3 36. I
ental formula: 1 == wa o0n nner upper
3-9’ © 7-7? PM3 9: Ma 5 pp
incisor about one-third higher than outer, and frequently bicuspidate ;
in western specimens this character varies greatly. Lower incisors
about equal in height and slightly crowded in the tooth-row; exceeded
in height by cingulum of canine. First upper unicuspid (pm?)
minute, the second (pm*) unusually broad, its greatest width
equaling its height from cingulum to tip. First and second lower
unicuspids (pm; and pm;) small, less than two-thirds height of third
(pm ;) ; all conical and with complete cingula. Cusps of lower molars
particularly high and sharp.
Limbs and Membranes——Wing short and broad, length of fifth
metacarpal equaling that of third; wing membrane attached at side
of metatarsus, just below bases of toes. Calcar neither keeled nor
340 University of California Publications in Zoology ([Vou.17
lobed, slightly less in length than free edge of interfemoral membrane.
Length of foot about half that of tibia. Tail exceeds forearm in
length by from 2 to 10 millimeters. All membranes relatively thin
and delicate.
Pelage.—F ur everywhere full and soft, longest on middle portion
of back, where it reaches a maximum length of 11 millimeters. Face
naked save for occasional long hairs rising from the glandular masses,
and a fringe of shorter finer hairs extending downward along sides
of upper lip. Fur reaches onto dorsal base of ear for a distance of
2 or 3 millimeters. Wings and interfemoral membrane naked except
at their extreme bases.
Corynorhinus rafinesquii pallescens Miller
Pale Lump-nosed Bat
Synotus Townsendii, Cooper (in Cronise, 1868, p. 442). Occurrence east
of the Sierras.
Synotus Townsendii, Cooper (1868, p. 6) (?).
Corynorhinus macrotis pallescens Miller (1897b, pp. 52-53, figs. 9-10).
Original description; type locality, Keam Canon, Arizona.
Corynorhinus macrotis pallescens, Elliot (1901, pp. 399-400). Diagnosis;
distribution.
Corynorhinus macrotis pallescens, Elliot (1904b, p. 604). Diagnosis; distri-
bution.
Corynorhinus macrotis pallescens, Elliot (1905, p. 491). Distribution.
Corynorhinus macrotis pallescens, Stephens (1906, p. 265). Diagnosis;
distribution ; habits.
Corynorhinus macrotis pallescens, Miller (1912, p. 67). General range.
Corynorhinus macrotis pallescens, Grinnell and Swarth (1913, pp. 379-380).
Oceurrence in San Jacinto region; habits.
Corynorhinus macrotis pallescens, J. Grinneil (1913b, p. 281). Range in
California.
Corynorhinus macrotis pallescens, J. Grinnell (1914, p. 263). Occurrence
at Riverside Mountain, Colorado River.
Corynorhinus macrotis pallescens, H. W. Grinnell (1914, p. 320). Com-
parison with C. m. intermedius.
Corynorhinus megalotis pallescens, G. M. Allen (1916, pp. 341-344), part.
Description; status of subspecies; list of localities.
Diagnosis —Color wood brown; total length, 92 to 101 millimeters,
averaging 97.2.
Description: Color—Dorsal surface wood brown, becoming paler
about head; hairs with faintly defined light plumbeous bases. Ventral
surface vinaceous buff; membranes a slightly darker shade of same
color. G. M. Allen (1916, p. 342) discusses color variation in C. r.
pallescens and records minutely the shading of two ‘‘reddish’’ indi-
viduals (10694, 10695, Mus. Vert. Zool.) secured in a mine at River-
side Mountain on the Colorado desert. J. Grinnell had earlier stated
(1914, p. 263) that the reddish east of these two specimens ‘‘is doubt-
less wholly adventitious, due to the fine, sticky red dust with which
the walls of the mine were covered.”’
1918] Grinnell: A Synopsis of the Bats of California 341
A specimen in the Museum of Comparative Zoology collected in
the hills back of Lone Pine is stated by G. M. Allen (loc. cit.) to be
““ pale pinkish buff’ above and nearly white below to the roots of
the hairs.”’
Measurements —Average and extreme measurements in milli-
meters of a series of ten specimens from the San Jacinto Mountain
region are as follows: total length, 97.2 (extremes, 92.0-101.0) ;
tail vertebrae, 47.7 (48.0-50.0); tibia, 18.1 (15.9-19.3); foot, 9.0
(8.0-11.0) ; forearm, 39.9 (37.5-41.8) ; greatest length of skull, 15.9
(15.5-16.4).
MEASUREMENTS IN MILLIMETERS OF TEN SPECIMENS OF CORYNORHINUS RAFINESQUII
PALLESCENS MILLER FROM NEAR KENWORTHY, SAN JACINTO
MounTAIN REGION, CALIFORNIA
A ae
ane: 5g Ge ys 5
a ae . $= as 23 55
1883 fot 97.0 46.0 18.0 8.0 39.3 16.0 8.3 8.8 3.4
1885 Jb 92.0 46.0 15) 8.0 39.3 1525 8.2 8.5 3.4
1887 ret 98.0 49.0 18.7 8.0 38.5 16.3 8.5 9.0 3.6
1888 Jb 93.0 46.0 15.9 8.0 37.5 16.4 8.2 8.8 y43)
1889 Jb 99.0 50.0 18.0 9.0 40.1 15.9 8.2 9.0 3.6
1890 3 93.0 43.0 18.0 11.0 41.7 15.8 8.0 8.9 3.5
1891 3 100.0 49.0 18.7 10.0 39.6 16.4 8.6 8.9 3.6
1892 fof 101.0 50.0 18.1 10.0 39.8 15.8 8.3 8.9 3.5
1884 io) 98.0 48.0 19.0 8.0 41.8 16.1 8.4 8.8 3.0
1893 2 101.0 50.0 19.3 10.0 41.0 15.9 8.2 9.0 3.4
Synonymy and History—Corynorhinus macrotis [== rafinesquit|
pallescens was described by Miller (1897b, pp. 52-53) from specimens
captured in Keam Cafion, Navajo County, Arizona. This author
regarded pallescens as a race of the Corynorhinus macrotis described
by LeConte (1831, p. 481). LeConte does not designate a type local-
ity, but Miller (1897), p. 51) fixes the type locality as the LeConte
Plantation, in Georgia. G. M. Allen (1916, pp. 838-341) in a revision
of the genus Corynorhinus states that the species of Corynorhinus
occurring west of the Alleghanies is distinct from the one described
by LeConte, and revives the name megalotis of Rafinesque for the
western animal. Later, Oldfield Thomas (1916, p. 127) finds the
name megalotis preoccupied, and to have been replaced with the
name rafinesquii by Lesson (1827, p. 96).
I have examined certain of the specimens in the United States
National Museum upon which G. M. Allen based his opinion, and
coneur with him in recognizing the distinctness of the two species,
and the nearer affinity of Californian Corynorhinus with the form
342 University of California Publications in Zoology (Vou. 17
rafinesquu. G. M. Allen (loc. cit.) fails to recognize the race inter-
medius, placing some individuals so called by the present writer
with the race pallescens and others with townsendii. This matter is
discussed on page 347 of the present paper.
|
re
AS
au @ CORYNORHINUS RAFINESQUIT
‘) PALLESCENS
& CORYNORHINUS RAFINESQUIL
INTERMEDIUS
! id Bema
RT AAO ex id.
a \ : Ce apes get
x) ou URADer S
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALLFORNIA
Fig. T. Map showing stations of occurrence in California of Corynorhinus
rafinesquit pallescens and Corynorhinus rafinesquii intermedius, as established by
specimens examined by the author.
Distribution—tThe distribution of this bat has not been fully
worked out, but indications are that it will be found to oceur through-
out the arid portions of the southwestern United States and extreme ~
northern Mexico (see G. M. Allen, 1916). In California it has been
found in the Lower and Upper Sonoran zones, chiefly on the Colorado
and Mohave deserts. (See map, text-fig. T.)
2 |
1918} Grinnell: A Synopsis of the Bats of California 343
Specimens Examined.—Total number 28, from the following
localities in California: Imperial County: Palo Verde, 1; San Diego
County: Julian, 1; Vallecito, 2; San Diego, 1 (San Diego Soc. Nat.
Hist.) ; Riverside County: near Kenworthy, San Jacinto region, 17;
Riverside Mountain, near Colorado River, 2; Whitewater, 1; San Ber-
nardino County: Oro Grande, 2 (U.S. Biol. Surv.) ; Inyo County:
Lone Pine, 1 (Mus. Comp. Zool.).
Natural History.—Stephens (1906, p. 265) states that the pale
lump-nosed bat appears to inhabit caves. All specimens of this bat
which have been taken in California have been, as far as the writer is
aware, captured in caves or mine tunnels, with the exception of those
secured by Stephens at Vallecito, San Diego County. This collector
writes (MS) that at Vallecito there is an old adobe house, vacant and
without windows or doors. When camped in the vicinity Stephens
made a practice of filling the windows with brush and hanging
blankets above the doorways, ready to be let down. After dark a
lantern was taken into the building and the blankets let down. As
the bats flew about they were secured with a butterfly net. They
did not come into the building until some time after dark, often an
hour after the last rays of daylight. The only bats secured in the
above manner were of the genera Corynorhinus and Macrotus.
The seventeen specimens taken May 22 and June 5, 1908, at Ken-
worthy, San Jacinto Mountains, were secured in an old mine tunnel
some 600 feet in leneth and were found scattered along singly, cling-
ing to the sidewalls from a few feet within the entrance nearly to
the end. The bats were cold and searcely able to move. The ears
were folded down the sides and almost completely hidden by the
wings which were held together in front, that is, against the rock. <A
female among the bats taken here contained one large embryo (see
Grinnell and Swarth, 1913, pp. 379-3880). Of these seventeen speci-
mens fifteen were males.
J. Grinnell (1914, p. 263) reports the finding on March 18, 1900,
of three bats of this species clinging to the rock wall at the end of
the sloping drift in the Steece copper mine at Riverside Mountain, on
the Colorado Desert. Both of the specimens secured were females
and one contained a single embryo.
Because of its fondness for caves and tunnels this bat is well
known to miners, who, in recognition of its long ears eall it the
““burro bat.’’
On March 15, 1915, a live male Corynorhinus r. pallescens was
344 University of California Publications in Zoology (Vou. 17
sent to the Museum of Vertebrate Zoology by Mr. W. C. Hanna,
who found it on March 13, in a cave near Whitewater, Riverside
County. When received at the Museum the bat was cold and torpid
and the long ears were folded backward and downward and against
the sides of the head; the long narrow tragi, however, stood rigidly
erect (pl. 17, fig. 12). The ears were not bent sharply back, but
curved by a regular and even crimping of the folds along the inner
edges of the auricles, the effect thus produced reminding one strongly
of the curving horns of a mountain sheep (pl. 15, fig. 4).
When the bat was taken into a warm room and handled his ears
gradually unbent (pl. 15, fig. 5) until they stood erect and directed
slightly forward (pl. 15, fig. 6). He soon became lively and protested
in shrill cicada-like notes, and when released he quickly took flight.
The aerial motions of the little mammal were fascinating to watch
as it repeatedly circled the room in its search for an exit. At one
moment it was drifting as easily and slowly as a butterfly on a sum-
mer breeze, at another it was skimming as swiftly as a swallow, down
almost to the surface of the floor, then up among the recesses of the
beamed ceiling, an instant hovering where a draft of air came
through the crack beside the closed door, then over to each window
in turn, hovering up and down before the pane, touching nothing but
the light cord which hung from the edge of each window-shade, and
which retreated before the fanning wings.
A wild bird loosed in a room stirs one to instant pity by its head-
long dash against the window glass; but the wild bat aroused only
keenest admiration, for the dexterity of his searching flight was such
that not a hair was ruffled.
Tired by his long flight the bat at last retreated to a beam of the
ceiling where he hung in a shaded recess, head down, but ears and
eyes alert. Finding himself unmolested his vigilance gradually re-
laxed, his eyes closed, his ears curled backwards, the tip of his tail
curled under him, his thumbs turned back under his wings, the tiny
claws thus aiding in the supporting of his body, and he was asleep.
Corynorhinus rafinesquii intermedius H. W. Grinnell
Intermediate Lump-nosed Bat
Corynorhinus macrotis townsendii, J. Grinnell (1913b, p. 281). Record
from Placer County.
Corynorhinus macrotis intermedius H. W. Grinnell (1914, p. 320). Orig-
inal description; type from Auburn, Placer County.
1918] Grinnell: A Synopsis of the Bats of California 345
Corynorhinus megalotis pallescens, G. M. Allen (1916, pp. 341-344), part.
Description; localities of record.
Corynorhinus megalotis townsendii, G. M. Allen (1916, pp. 344-347), part.
Description; localities of record.
Diagnosis—Similar in general characters to Corynorhinus rafi-
nesquiu pallescens Miller and Corynorhinus rafinesqui townsendit
(Cooper), but intermediate in color between these two forms; darker
colored and larger than pallescens.
Description—aAs compared with pallescens, intermedius is some-
what larger in general size; ten examples of the latter form from
Auburn, Placer County, average 102 millimeters in total length, while
ten specimens of pallescens from the San Jacinto region average but
97.2 in the same dimension. In color intermedius is natal brown
above; below, wood brown; membranes bone brown.
Measurements——A series of ten specimens from west-central Cali-
fornia shows averages and extremes in millimeters as follows: Total
length, 102 (extremes, 97.0-108.0) ; tail vertebrae, 48.6 (45.0—-55.5) ;
tibia, 19.7 (18.7-21.0); foot, 9.8 (9.0-12.5); forearm, 42.0 (40.2—
43.6); greatest length of skull, 16.2 (15.4-17.1).
Distribution —tThe evidence at hand indicates that this bat oceu-
ples a geographic position intermediate between that of pallescens
and that of townsendii, namely the semi-arid and semi-humid portions
of the Upper Sonoran zone in California west of the desert divides.
(See map, text-fig. T.)
Specimens Examined.—Total number, 44, from the following
loealities in California: Tulare: County: Eclipse Mine, White River,
12 (Stanford Univ.) ; Placer County: Auburn, 23; Pioneer Cave, 3;
Santa Catalina Island: Johnson Harbor, 1; Napa or Sonoma County :
Mount Veeder, 1 (U.S. Biol. Surv.) ; Siskiyou County: Happy Camp,
1 (U. S. Biol. Surv.) ; San Benito County; Bear Valley, 2 (U. S.
Biol. Surv.) ; Hernandez, 1 (Calif. Acad. Sci.).
Natural History —tThere are in the Museum of Vertebrate Zoology
three females of intermedius taken in Pioneer Cave, Placer County,
May 12, 1870, by J. G. Cooper; and five females and one male taken
at Auburn, Placer County, July 31, 1909, by Dr. J. C. Hawver. On
August 6, 1913, Dr. Hawver secured fifteen females and two males
in the belfry of the grammar school at Auburn. Dr. Hawver stated
in a letter that these bats oceur at Auburn the year around, showing
no signs of being migratory.
Remarks—An example of intermedius from Mount Veeder, on
the border between Napa and Sonoma counties, varies strongly in the
direction of townsendii. While Mount Veeder is not exactly within
the humid coast belt, yet it lies in a region where there is a strong
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1918 | Grinnell: A Synopsis of the Bats of California 347
infusion of species belonging typically to the fauna of the humid
coast district.
According to Miller (1897), p. 53) townsendw is the form of
Corynorhinus macrotis [= rafinesquii] occupying the humid coast
belt of Oregon, Washington and British Columbia. No examples of
any form of Corynorhinus which may occupy the humid coast district
of northern California are available and it is entirely possible that
the bats from this area may prove to be true townsendi.
G. M. Allen (1916, pp. 343-347) fails, in his revision of the genus
Corynorhinus, to recognize the race intermedius and divides the type
series between the races pallescens and
townsendii. Of the nine skins of inter- \
medius from the type locality which he ~~ /
examined, he places two as pallescens,
“not typieal,’’ and the remaining seven
as townsendii, ‘‘not all typical.’’ <A
skin of intermedius from Catalina
Island is listed as pallescens, ‘‘not typi-
”
eal. I have carefully reéxamined the Fic. U
g. U.
y Front view of head
series of skins upon which the name of Antrozous pacificus, X 1.00,
: - é showing nostrils opening for-
intermedius was based, and again com- ward beneath horseshoe-shaped
ridges, long entirely separated
4 é 4 ears, and long tapering tragi,
townsendii; but even with the aid of with slightly crenulate posterior
borders.
pared it with series of pallescens and
Dr. Allen’s paper I am unable to relin-
quish my opinion that specimens from the semi-humid and semi-arid
portions of west-central California constitute a recognizable race,
intermedius. It would indeed be extraordinary if two subspecies
should have existed in a single colony !
Subfamily NycroPHILINAE
This subfamily, which is represented in North America by but a
single genus, differs from the Vespertilioninae in the abruptly trun-
cate muzzle, on the anterior face of which the nostrils open forward
beneath a distinct horseshoe-shaped ridge or small nose-leaf (Miller,
1907, p. 284).
Genus Antrozous H. Allen
Ears separate (text-fig. U), extending considerably beyond tip
of muzzle when laid forward; tragus slender, straight, tapering, a
348 University of California Publications in Zoology (Vou. 17
little less than half as high as ear conch; terminal joint of tail ex-
serted. (For skull and teeth, see under species. )
The genus as occurring in California is represented by two closely
related species, Antrozous pallidus and A. pacificus.
Antrozous pallidus (Le Conte)
Desert Pallid Bat
V [espertilio|. pallidus LeConte (1855, p. 437). Original description; type
locality, El Paso, Texas (Baird, 1859, II, p. 5).
Vespertilio pallidus, Baird (1859, II, pp. 4-5), part. Description; re-
corded from ‘‘ California. ’’
Antrozous pallidus, H. Allen (1864, pp. 68-69, 3 figs. in text), part. De-
scription; distribution (includes ‘‘Ft. Yuma’’).
Antrozous pallidus, Coues (1867, p. 283). Habits.
Antrozous pallidus, Cooper (1868, p. 6), part. Distribution.
Antrozous pallidus, Cooper (in Cronise, 1868, p. 442). Occurrence in Cali-
fornia.
Antrozous pallidus, Coues and Yarrow (1875, pp. 85-86), part. Diagnosis;
range; habits:
Antrozous pallidus, Dobson (1878, p. 171, pl. 11, figs. 6-6b). Description.
Recorded from ‘‘California’’ and ‘‘Mammoth Valley, California.’’
Antrozous pallidus, Bryant (1891a, p. 358), part. Nominal.
Antrozous pallidus, H. Allen (1894, pp. 66-70, pl. 8), part. Description;
distribution.
Antrozous pallidus pallidus, Miller (1897b, pp. 43-45, pl. 1, fig. 10, 4 figs.
in text). Description; distribution; recorded from Fort Yuma, Owens
Valley, Panamint Valley, and Walker Basin.
Antrozous pallidus, Elliot (1901, pp. 396-897, fig. 82). Description; general
distribution.
Antrozous pallidus, Miller and Rehn (1901, pp. 266-267). Type locality.
Antrozous pallidus, Elliot (1904a, p. 319). Distribution in California;
recorded from Lone Pine and Coso Mountains, in Inyo County.
Antrozous pallidus, Elliot (1904b, pp. 605-607, fig. 115). Description;
general distribution.
Antrozous pallidus, Stephens (1906, p. 263, fig. in text). Description;
distribution.
Antrozous pallidus, Elliot (1907, p. 518). Records repeated.
» Antrozous pallidus pallidus, Lyon and Osgood (1909, pp. 278-279). Loca-
tion and condition of type.
Antrozous pallidus paliidus, Miller (1912, p. 68). General range.
Antrozous pallidus pallidus, J. Grinnell (1913b, p. 282). Range in Cali-
fornia.
Antrozous pallidus pallidus, J. Grinnell (1914, p. 263). Occurrence along
Colorado River.
Diagnosis —F orearm 46 to 51.4 millimeters long; greatest length
of skull, 19.4 to 21.0; color, whitish drab gray.
Description: Head.—Muzzle squarely truneate, with a low but dis-
—
1918] Grinnell: A Synopsis of the Bats of California 349
tinct horseshoe-shaped ridge above each nostril (text-fig. U) ; behind
this, a large flattish swelling on each side. Ears separate and wide
apart at bases, extending considerably beyond tip of muzzle when laid
forward (pl. 17, fig. 11). Tragus slender, straight and long, half
leneth of ear. Whole posterior border of tragus faintly crenulate; a
well-developed lobe at posterior base.
Limbs and Membranes.—Wing short and broad; third metacarpal
but slightly longer than fifth. Lateral membrane attached at bases
of toes. Tip of tail exserted beyond interfemoral membrane. Length
of calear somewhat less than half that of free border of interfemoral
membrane. Feet broad and strong, about half as long as tibia.
Claws large and strong; chord of exposed portion of each claw about
2.5 millimeters long.
Pelage—¥F ur on back silky, and about 8 millimeters in length.
Fur on ventral surface shorter, averaging about 5 millimeters in
leneth, and more abundant. Wing and tail membranes not furred
except at their extreme bases. A narrow strip of hairs on dorsal
surface of anterior border of ear, extending outward to about middle
of border, and diminishing in width as it advances. On ventral sur-
face of ear, lines of hair extend along two ridges which run parallel
with anterior border of auricle. A few short hairs on backs of toes.
Color.—Hairs on back very pale drab gray, tipped on terminal
fourth with pale brown. On under surface of body the hairs are
erayish white, and toward sides of body suffused with a pale yellowish
tint. Ears light brownish, wing membranes and feet darker.
Skull—Greatest length varies from 19.4 to 21.0 millimeters.
Brain-case, rostrum, and palate, broad. Length of bony palate
behind molars (exclusive of median spine) slightly less than its width
at base of median spine. Dorsal profile unevenly convex. Rostrum
more than half as long as brain-ease. Auditory bullae covering nearly
the entire cochleae; their greatest diameter equal to twice the distance
between them. (See pl. 23, fig. 55; pl. 24, fig. 63.)
Teeth—Dental formula: 1 ea) at pm = m = —— 8 Upper
incisor large and simple, its shaft more than half as high as canine.
Posterior basal lobe of incisor in contact with cingulum of canine.
Lower incisors somewhat crowded; their trilobed crowns strongly
imbricated. »- Upper and lower canines about equal in length, but
former twice as wide at base as latter; cingula of both distinct but
small. Upper premolar transversely long and narrow. First lower
premolar small and narrow, closely wedged between canine and
second premolar; latter equals first lower molar in height. First and
second upper molars with middle cones highest. Third upper molar
with less than half crown area of either of others, its outer cone
highest. Lower molars with outer cusps much higher than inner.
Measurements—Average and extreme measurements in milli-
meters of a series of eight specimens in the Museum of Vertebrate
Zoology are as follows: Three males, total length, 104.0 (extremes,
99.0-111.0) ; tail vertebrae, 42.3 (40.0-47.0) ; tibia, 18.9 (18.4-19.4) ;
foot, 11.8 (11.0-12.0) ; forearm, 47.4 (46.8-48.4) ; greatest length of
skull, 19.7 (19.4-20.2) ; five females, total length, 112.3 (103.0-122.0) ;
350 University of California Publications in Zoology [Vou.17
tail vertebrae, 47.0 (41.0-53.0) ; tibia, 20.1 (19.0-21.0); foot, 11.8
(11.5-13.0) ; forearm, 50.9 (50.6-51.4) ; greatest length of skull, 20.6
(20.4-21.0).
MEASUREMENTS IN MILLIMETERS OF NINE SPECIMENS OF ANTROZOUS PALLIDUS
(LEConTR), FROM CALIFORNIA
oe
a ny ° Sa BS
Fs 2 E gee 98 32 22
Mus. ge 5 5 =) z ga =e 32 BE
no. Sex. 57 a” - & i ee hia pat ale
5837* ot) Rede 0) yeceed 2 Ope SII. ees i
73407 ie) 122.0 53.0 21.0 13.0 50.8 21.0 12.5 10.0 4.2
73417 io) 113.0 46.0 20.6 12:0 ese 20.6 12.2 O27; 4.0
7342? 2 103.0 41.0 20.0 11.5 51.4 20.6 12.2 9.4 3.9
73437 3 102.0 40.0 19.0 11.0 47.2 19.6 11.5 9.0 4.0
73447 fof 99.0 40.0 18.4 11.0 46.8 19.4 11.6 9.2 3.9
73457 OF 7 A1320 45.0 19.0 WEST, See 20.4 12.0 9.7 3.9
T3462) ge - dE0 47.0 19.4 12.0 48.4 20.2 11.8 9.3 4.1
10696° 2) 112.0 50.0 20.3 12.0 50.6 20.4 11.4 9.5 3.8
1 From Swansea. Inyo County.
*From Vallecito, San Diego County.
*From Colorado River, Imperial County.
Synonymy and History.—Antrozous pallidus was first deseribed
by LeConte (1855, p. 487) under the name Vespertilio pallidus. In
the original description the type locality is given as California, but
both Baird (1859, II, p. 5) and H. Allen (1864, p. 69) have stated
that the type material came from El Paso, Texas. The type is now
in the United States National Museum and according to Lyon and
Osgood (1909, pp. 278-279) is in a good state of preservation and
clearly referable to the desert form.
H. Allen in 1862 (p. 247) made pallidus the type species of the
new genus Antrozous.
Merriam in 1897 (p. 179) deseribed the race of Antrozous from
the region west of the deserts under the name Antrozous pallidus
pacificus, showing it to be distinet from the form deseribed by Baird.
Distribution.—The range of A. pallidus is given by Miller (1912,
p. 68) as being the Lower Austral zone in the desert region of eastern
California, Nevada, Arizona, New Mexico, and western Texas. War-
ren (1910, p. 284) gives two records for Colorado. In California
this species occupies the Lower Sonoran zone on the Colorado and
Mohave deserts. (See map, text-fig. V.)
Specimens Examined.—The writer has examined 9 specimens of
this bat from the following localities in California: Imperial County :
1918] Grinnell: A Synopsis of the Bats of California 351
Colorado River, 8 miles east of Picacho, 1; San Diego County: Valle-
cito, 7; Inyo County: Swansea, 1.
Natural History.—tThe earliest account of the habits of this species
with which the writer is familiar is that given by Coues (1867, pp.
ANTROZOUS PALLIDUS
ANTROZOUS PACIFICUS
ANTROZOUS (PACIFIOUS?) |
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OF CALIFORNIA s
Fig. V. Map showing stations of occurrence in California of Antrozous pal-
lidus and Antrozous pacificus, as established by specimens examined by the author.
283-284). This author found the desert pallid bat very abundant
at Fort Yuma, where during the hot months it became a decided
nuisance. The proximity of the crowded retreats of the bats in the
chinks and crannies of the officers’ quarters was actually offensive.
At night the bats fluttered about the rooms by scores, and by day they
352 University of California Publications in Zoology |Vou.17
could be continually heard scratching and squeaking in their retreats.
When caught or disabled they squeaked harshly and bit with vigor
and considerable effect, if incautiously handled.
Bailey (1905, pp. 214-215) reports these bats as occurring in west-
ern Texas between April 18 and October 11, but adds that these limits
are apparently dates of collectors’ entering and leaving the region
rather than of the migration of the bats. During the day the bats
were found by the collectors in crevices of buildings and behind sign-
boards, and, as they occurred also in rocky country where there were
no buildings, Bailey considers it probable that they there hid in
crevices of the cliffs during the day. This author describes their
flight as soft and noiseless, and while rapid, not so jerky and quick
as that of most bats. He adds that the light color and large size
render this species of bat unmistakable in the early evening.
J. Grinnell (1914, p. 263) states that a female taken April 20
on the Colorado River, eight miles east of Picacho, contained two
embryos; and Stephens (MS) notes that a specimen taken at San
Bernardino, May 16, contained three embryos.
Antrozous pacificus Merriam
Pacifie Pallid Bat
Vespertilio pallidus, Baird (1859, II, pp. 4-5), part? Description; recorded
from ‘‘California.’’
Antrozous pallidus, H. Allen (1864, pp. 68-69, 3 figs. in text), part. De-
scription; recorded from ‘‘Posa Creek,’’ Tejon Valley, and Ft. Tejon,
Kern County.
Antrozous pallidus, Cooper (1868, p. 6), part. Distribution.
Antrozous pallidus, Coues and Yarrow (1875, pp. 85-86), part. Diagnosis;
range; habits.
Antrozous pallidus, H. Allen (1894, pp. 66-70, pls. 8-9), part. Description.
Antrozous pallidus pacificus Merriam (1897, pp. 179-180). Original de-
scription; type locality, Fort Tejon.
Antrozous pallidus pacificus, Miller (1897b, pp. 45-46, 1 fig. in text). De-
scription; distribution; several California localities listed.
Antrozous pallidus pacificus, Elliot (1901, p. 397). Diagnosis.
Antrozous pallidus pacificus, Miller and Rehn (1901, p. 267). Type
locality.
Antrozous pallidus pacificus, Elliot (1904a, pp. 319-320). Occurrence at
Fort Tejon; habits.
Antrozous pallidus pacificus, Elliot (1904b, p. 607). Diagnosis; distri-
bution.
Antrozous pallidus pacificus, Stone (1904b, p. 587). Record of specimens
secured at Berkeley.
Antrozous pallidus pacificus, Stephens (1906, p. 263). Diagnosis; distri-
bution.
1918] Grinnell: A Synopsis of the Bats of California 353
Antrozous pallidus pacificus, Elliot (1907, p. 518). Record of specimens
taken at Fort Tejon and San Rafael.
Antrozous pallidus pacificus, Lyon and Osgood (1909, pp. 278-279). Loca-
tion and condition of type.
Antrozous pallidus pacificus, Miller (1912, p. 68). General range.
Antrozous pallidus pacificus, J. Grinnell (1913b, p. 282). Range in Cali-
fornia.
Diagnosis —Forearm 53.7 to 58.9 millimeters; greatest length of
skull, 22.0 to 23.9; color, yellowish drab brown.
Description—In general characters Antrozows pacificus is very
similar to Antrozous pallidus, but is slightly larger, with decidedly
larger skull (pl. 23, fig. 53, and pl. 24, fig. 61), and darker coloration.
Limbs, membranes, and pelage essentially similar to those of pallidus.
Color—On the upper surface the hairs are pale yellowish drab
at their bases, sepia or drab on distal third. The light bases of the
hairs show in irregular patches. According to Stephens (1906.
p. 263) there is a patch on the back of the neck, and sometimes one
on the rump, where the dark tips are lacking. This is difficult to
determine in some of the dried skins. On the under surface the hairs
are cream color, shghtly darker at the tips.
Skull—tThe skull varies in greatest length from 22 to 23.9 milli-
meters. The brain-case, rostrum and bony palate are considerably
broader than in pallidus. The length of the bony palate behind the
molars (exclusive of the median spine) is usually equal to or greater
than width at base of median spine.
Teeth—tThe teeth are larger than those of pallidus, but similar
in form, save that the upper premolar is conspicuously broader and
shorter.
Measurements.—Average and extreme measurements of a series
of seventeen specimens in the Museum of Vertebrate Zoology are as
follows: ten males: total length, 118.0 (extremes, 109.0-125.0) ;
tail vertebrae, 44.5 (39.0-49.0) ; tibia, 19.9 (19.0-22.0); foot, 13.1
(11.0-16.0) ; forearm, 55.0 (54.0-56.0) ; greatest length of skull, 22.4
(22.0-23.0) ; seven females: total length, 118.8 (114.0-122.0) ; tail
vertebrae, 43.4 (41.0-46.0) ; tibia, 19.9 (18.0-21.7) ; foot, 13.4 (18.0-
16.0); forearm, 55.3 (53.5-58.9) ; greatest length of cranium, 22.6
(22.0-23.9).
Synonymy and History.—H. Allen (1864, p. 68) remarked that
in the species (Antrozous pallidus) two varieties of color are observed,
but he does not attempt to separate the possible forms indicated.
Many years later, C. H. Merriam (1897, pp. 179-180) described the
darker race under the name pacificus, selecting his type from three
specimens secured at Old Fort Tejon, California. The uniformly
much larger skull of pacificus, together with lack of imtergradation
with pallidus, causes the present writer to consider pacificus a distinct
species, rather than a race of pallidus.
Distribution —The general range of A. pacificus is given by Miller
(1912, p. 68) as ‘‘Austral zones of the western United States and
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1918] Grinnell: A Synopsis of the Bats of California 355
northwestern Mexico.’’ In California the range is through the Lower
and Upper Sonoran zones on the Pacific slope, from the Mexican line
north through the San Diegan district and San Joaquin Valley, and
through the central coast district as far as Marin County. Recorded
also from Fort Crook (near Burgettville), Shasta County (Miller,
1897b, p. 45). The specimen from Goose Lake, Modoc County, exam-
ided by me differs somewhat from other specimens of Antrozous which
I have seen and suggests the existence in the northern part of the
the Great Basin of a race, as yet unnamed, externally most nearly like
pacificus. (See map, text-fig V.)
Specimens Examined.—The writer has examined 107 specimens of
the Pacific pallid bat from the following localities in California: San
Diego County: Julian, 1; Los Angeles County: Glendora, 7 (U.S.
Nation. Mus., 1; Mus. Vert. Zool., 6) ; Pasadena, 12; Sierra Madre, 3:
Alhambra, 1 (U. 8. Nation. Mus.) ; Ventura County: Santa Paula, 15
(Stanford Univ.) ; Kern County: Fort Tejon, 2 (U.S. Nation. Mus.) ;
Kelso Valley, 1; San Luis Obispo County: Carrizo Plains, 2; Tulare
County: White River, 1 (Stanford Univ.) ; Fresno County: Fresno,
3 (U. S: Nation. Mus., 2; Mus. Vert. Zool., 1); Monterey County:
Carmel Mission, 2 (Stanford Univ.) ; Merced County: Snelling, 20;
Santa Clara County: Palo Alto, 19 (Stanford Univ., 17; Mus. Vert.
Zool., 2); Stanford University, 3 (Stanford Univ.) ; Mountain View,
1 (Stanford Univ.); San Mateo County: Menlo Park, 4 (Stanford
Univ.) ; Alameda County: Berkeley, 8 (U. C. Dept. Zool., 4; U. 8.
Nation. Mus., 1; Mus. Vert. Zool., 3); Marin County: San Geronimo,
1 (Stanford Univ.) ; Modoe County: Goose Lake, 1 (San Diego Soc.
Nat. Hist.).
Natural History—During the summer of 1904 the writer became
acquainted with a colony of the Pacifie pallid bats which occupied
the spaces under the eaves of a barn loft at Glendora, Los Angeles
County, in company with a few Mexican free-tailed bats. Every
morning the floor of the loft was found to be strewn with heads, legs,
and wings of insects caught by the bats. Most numerous among these
?
‘‘kitehen middens’’ were heads and legs of Jerusalem crickets and
the wings of sphinx moths. Dr. H. C. Bryant of the State Fish and
Game Commission has kindly identified a sample lot of these remains,
picked up in September, 1904, as belonging to the following genera
and species of insects: Prionus californicus, Stenopelmatus sp., Deile-
phila lineata, Microcentrum sp., Ligyrus gibbosus, and Gryllus sp.
It is of interest to note that the Jerusalem crickets (Stenopelmatus),
356 University of California Publications in Zoology [Vou.17
being wingless, must have been secured upon the ground, the bats
probably becoming aware of their whereabouts through hearing them
as they crawled about on the surface of the ground.
Of six specimens of this bat taken from this colony August 6, 1904,
four were females and two males, all adult.
Heller (in Elliot, 1904a, pp. 319-320) found this species rather
rare at the type locality, Fort Tejon. He says of it:
Several were secured while I was stopping in an old house. The bats could
not be found in the house during the day, but at night they entered through
the open windows, bringing with them large brown mole crickets, which they
devoured at their leisure while suspended from the roof. The floor of the
house below their perches was covered with the remains of the insects.
A female of this species (no. 14654) secured by Swarth and Carr
May 27, 1911, at Painted Rock, Carrizo Plains, San Luis Obispo
County, contained two embryos. Stephens (1906, p. 263) states
that the young of this bat are born about the first of July. Merriam
(1897, p. 180) found two females, taken June 28, 1891, at Fort Tejon,
to contain ‘‘large embryos, nearly ready for birth.”’
While earrying on field work at Snelling, Merced County, May
27, 1915, Mr. Charles L. Camp came upon a troop of youngsters who
were setting up a wire net under the gable end of a church building
with the intent of ridding the place of a colony of bats the presence
of which had caused annoyance to the pastor and his congregation.
Mr. Camp writes (MS):
I could plainly hear the bats squeaking (6:30 P.M.) inside the wall, at a
distance of fifty feet from the building. About 7:30 the first bat came out,
missed the net and flew away. It was quickly followed by another which circled,
lit on the side of the building, and was shot. No more bats appeared for ten
minutes and then they began to drop into the net at the rate of about four a
minute, coming faster toward the last, when twenty-one were found in the trap.
After the net was taken down at 8 o’clock, many more bats, all apparently of the
same species (Antrozous pacificus) flew out of the building, circled about in
the trees and flew away. The bats caught (22 in all) were females, each contain-
ing one or two embryos, except one which was a male with testes small or medium.
Bees and red-shafted flickers were living in the same building with
the bats.
With one exception all dates of occurrence known to me are be-
tween March 27 and October 19. In the collection at Stanford
University are two females secured by C. J. Pierson, January 1, 1895,
at Carmel Mission, Monterey County. The Pacific pallid bat is
probably to a partial extent migratory.
1918] Grinnell: A Synopsis of the Bats of California 357
Family MOLOSSIDAE
Foot and leg short and stout. Ears variable, sometimes joined
across forehead ; tragus much reduced; antitragus usually very large;
anterior border of auricle never with basal lobe. Muzzle obliquely
truneate, usually sprinkled with short modified hairs having spoon-
shaped tips; nostrils simple, usually opening on a special pad, the
upper surface of which is often set with fine horny excrescences.
Wing narrow; fifth metacarpal only about two-thirds as long as third;
membranes thick and leathery. Uropatagium short; tail projecting
conspicuously beyond its free edge. Miller (1907, p. 243) recognizes
ten genera, only two of which have been found to occur in the western
hemisphere north of southern Mexico.
Genus Nyctinomus Geoffroy
Ears large and rounded, rising from middle point of forehead,
extending distinctly beyond extremity of muzzle when laid forward;
anterior border of conch with 6 to 12 horny execrescences; keel well
developed; tragus small, flattened, squarely truncate above; anti-
tragus variable in form, sometimes low and indistinet. Muzzle pad
well developed and sharply outlined, its upper margin thickly set with
horny points like those on anterior margin of ear; a line of similar
points extends downward across middle of pad between nostrils.
Upper lip full and wrinkled, rather thickly sprinkled with stiffened
spoon-shaped hairs; feet stout; wings narrow (Miller, 1907, p. 251).
Nyctinomus depressus Ward
Tacubaya Free-tailed Bat
Nyctinomus depressus Ward (1891, pp. 747-750, 2 figs.). Original deserip-
tion; type locality, Tacubaya, Federal District, Mexico.
Nyctinomus macrotis nevadensis H. Allen (1894, pp. 171-174, pls. 34-35).
Deseription of supposed new form, from ‘‘ Nevada and California.’’
Nyctinomus nevadensis, J. A. Allen (1894, pp. 326-828). Description of
adult male; type fixed.
Nyctinomus nevadensis, Miller and Rehn (1901, p. 273). Type locality.
Nyctinomus macrotis nevadensis, Elliot (1901, p. 417). Description; dis-
tribution.
Nyctinomus nevadensis, Miller (1902, p. 250). Declared to be a synonym
of Nyctinomops depressus.
Nyctinomops depressus, Elliot (1904b, p. 627). Deseription; distribution.
Nyctinomops depressus, Stephens (1906, p. 275). Description; distribution.
Nyctinomus macrotis nevadensis, Lyon and Osgood (1909, p. 280). Nature
and location of type.
Nyctinomus depressus, J. Grinnell (1913b, p. 283). Nature of occurrence
in California.
Diagnosis —Size large; total length 131 millimeters, free portion
of tail, 33; muzzle not ‘‘concave’’ between ears; color, above, burnt
umber; below, Prout’s brown.
358 University of California Publications in Zoology [Vou.17
Description: Head—Inner edge of ear evenly convex when flat-
tened out; forward interior margin reflexed over a deep depression
at upper extremity of keel, forming a sort of pocket; outer margin
bilobate ; the lower lobe arising from a short straight base coming up
from antitragus, upper lobe continuous with tip and inner edge; keel
large, strongly reflexed at angle near base, and extending slightly
exterior to antitragus. Tragus straight on inner and upper margins;
outer margin formed by two shghtly concave lnes producing by their
juncture a slight lobe at center of this margin. Conch with seven
diagonally transverse flutings, appearing as furrows on upper surface,
and as ridges on lower surface, the posterior one being but very
-shehtly indicated. On outer surface of conch, and passing through
centers of these flutings at right angles to them, is a slight ridge
formed by a doubling of the skin. The peculiar depressed angle
formed by the juncture of the two lobes of the external margin of
the conch gives the ears of this species a peculiar drooping appearance
that suggested the specific name depressus. Nostrils circular, open-
ing forwards, outwards, and very slightly downwards. A prominent
subcireular swelling between eye and nostril and slightly below the
line connecting them. Side of face with five flutings extending to
lip, and a deep furrow under eye. Face and chin nearly naked.
Limbs and Membranes—On upper surface of antebrachial mem-
brane, a line of very short hairs, so fine as scarcely to be perceptible
when wet, borders humerus and radius. On upper surface of the
wing membrane, short lines of hairs border the radius, except at the
extreme elbow, and occupy the angle formed by juncture of fourth
and fifth metacarpals. Both upper and lower surfaces of wing mem-
brane covered with hairs to a line extending from proximal third of
humerus to middle of femur. Interfemoral membrane covered for
three or four millimeters below the femora on upper surface, but
naked on lower. <A well-developed ecallosity at base of first phalanx
of thumb.
Color—Above, burnt umber; below, Prout’s brown; bases of hairs
on both surfaces white. Membranes and ears (in aleohole specimen)
nearly concolor with under surface of body. Wing membranes, from
inner surface of distal end of tibia and from ealear, indefinitely
edged with yellowish white, this color better defined in centers of
interdigital spaces and in center of space between fifth finger and
tibia. Outer edges of first and fifth toes closely fringed with short
curved white hairs; on fifth toe, dorsal to this outer fringe, is a row
of less numerous curved hairs, exceeding others about three times
in length. From upper surface of base of each claw spring three or
four long curved hairs, about 8 millimeters in length of chord.
= ps 9-9 a
Teeth —Dental formula: = = ge join ===. eee
Day leah 2-2 3-3
incisors semi-conical, parallel, and separated by a space of 1 milli-
meter. Lower incisors bifid, crowded; the middle pair in a straight
line; the outer ones start from near centers of inner surfaces of middle
pair and diverge at an angle of 45 degrees from them. Canines long,
with distinct, unbroken cingula, somewhat dilated on posterior in-
ternal part of lower one, but not forming a true cusp. Upper canines
—30)) Upper
1918 | Grinnell: A Synopsis of the Bats of California 309"
eurved backwards, sabre-shaped, passing 1.5 millimeters below gums
of lower teeth when mouth is closed, lower pair fitting into sockets
between upper incisors and canines. First upper and lower pre-
molars much smaller than second ones. Second upper premolars
decidedly longer than molars, with very acute outer cusps; internal
cusps not particularly developed.
Measurements (in millimeters from alcoholic specimen) .—Length
of head and body, from tip of nose to base of tail, 79; length of tail,
52; length of tail beyond interfemoral membrane, 33; length of head,
31; length of ear, from notch between antitragus and coneh to an-
terior point of margin, 25; length of antitragus, 7; height of anti-
tragus, 4.5; width of tragus at top, 2.5; ears united at base for, 3.5;
leneth of forearm, 60; length of thumb, not including metacarpal, 8;
extent of outstretched wings, 357; length of tibia, 18; leneth of foot,
13; length of caleaneum (poorly defined), about 16.
The above description is compiled from Ward’s (1891, pp. 747—
750, original description). A comparison of this deseription with
those given by H. Allen (1894, p. 171) and J. A. Allen (1894, p. 326)
shows agreement in general characters. Ward, however, notes the
absence of a gular pouch, while J. A. Allen indicates the presence of
a well-marked gular pouch.
Synonymy and History—This bat was described by Ward (1891,
pp. 747-750) from a single adult male specimen, taken within the
museum building at Tacubaya, Federal District, Mexico. In 1893,
H. Allen (1894, pp. 171-174, pls. 34, 35), apparently unaware of
Ward’s description, redescribed the species as Nyctinomus macrotis
nevadensis, from two immature specimens in the United States
National Museum. J. A. Allen (1894, pp. 326-328), upon examin-
ation of a fully adult male, raised the supposed form to full specific
Lolisy =
rank, and fixed typeship upon a specimen (no. 365607 U. S. Nation.
odd00E
Mus.) from ‘‘California.’”’ Miller (1902, p. 250) placed the name
nevadensis of H. Allen under the synonymy of Nyctinomus depressus
Ward.
Distribution—tThis bat has been recorded from Mexico City,
Mexico, and within the United States from Arizona, Nevada, Colo-
rado, Iowa, and California, there being but one record for each of
the states named, except for Iowa, where two specimens have been
secured (Cory, 1912, p. 477; Gabrielson, 1916, p. 86). :
In the cases of the Nevada and California examples of Nyctinomus
depressus the exact localities of capture are not known. The Colo-
rado example was taken at Grand Junction (altitude about 5000
feet) (Warren, 1910, p. 286) ; and the Arizona specimen when secured
was flying over a small meadow in the Chiricahua Mountains, at an
altitude of about 9500 feet (J. A. Allen, 1895, pp. 245-246). Mexico
360 University of California Publications in Zoology (Vou. 17
City, where the type specimen was taken, has an altitude of over
7000 feet.
Natural History.—Nothing is known under this head.
Nyctinomus femorosaccus Merriam
Pocketed Bat
Nyctinomus femorosaccus Merriam (1889a, p. 23). Original description;
type locality, Agua Caliente [= Palm Springs, Riverside County], Cali-
fornia.
Nuyuctinomus femorosaccus, Bryant (1891a, p. 359). Nominal.
Nyctinomus femorosaccus, Bryant (1892, p. 220). Nominal.
Nyctinomus femorosaccus, Elliot (1901, pp. 417-418). Description.
Nyctinomus femorosacecus, Miller and Rehn (1901, p. 272). Type locality.
Nyctinomus femorosaccus, Elliot (1904a, p. 321). Specimen recorded from
Palm Cafion, Colorado Desert.
Nyctinomus femorosaccus, Elliot (1904b, p. 626). Description.
Nyctinomops femorosaccus, Stephens (1906, p. 274). Description; type
locality.
Nyctinomops femorosaccus, Elliot (1907, p. 523). Repeated record.
Nyctinomus femorosaccus, Miller (1912, p. 69). Nominal.
Nyctinomus femorosaccus, J. Grinnell (1913), p. 282). Range in Cali-
fornia.
Diagnosis —Size medium (total length 103 millimeters) ; tail more
than half exserted; a fold of membrane extends from inner third of
femur to middle of tibia, forming a pocket at thigh; ears connected
at bases; color dull brown.
Description.—Merriam (1889a, p. 23) gives the following deserip-
tion of the type specimen of the pocketed bat:
Measurements [in millimeters] (from the alcoholic specimen).—Total length,
103; head and body, 60; tail, 41; exserted part of tail, 23; head, 23; ear from
crown, 14; ear from base of antitragus, 20; tragus, 1; humerus, 28; forearm,
47; third finger: metacarpal, 45; first phalanx, 20; second phalanx, 19; fifth
finger, 44.
General Characters.—Incisors = Lower incisors bifid and crowded; first
upper premolar small, but well developed; second very large, with a large and
high antero-internal cusp. Ears thick, united by bases of inner margins 4.5 mm.
from end of nose; ear keel greatly developed, with a large lobe on its lower third;
antitragus higher than long, convex anteriorly, slightly concave posteriorly, and
separated by a deep notch; tragus subquadrate, hidden behind the large anti-
tragus, its outer angle projecting upward in the form of a small pointed lobule;
upper margin of ear conch with two minute horny projections, not symmetrical
on the two sides. Tail more than half exserted. Gular sae present (opening on
right side of median line). There is a curious fold of membrane stretching from
the inner third of the femur to the middle of the tibia, forming a deep pocket
between it and the interfemoral membrane. The wing membrane is attached
to the leg at the same point (immediately below the middle of the tibia), so
that there are three folds of membrane here. The fur extends out on the wing
membrane, above and beneath, as far as a line drawn from the middle of the
humerus to the junction of the middle and outer thirds of the femur. Color,
dull brown. c
1918 | Grinnell: A Synopsis of the Bats of California 361
36038
20922
Huachuea, Arizona, which the present writer has examined, measures,
in millimeters: total length, 19.2; zygomatie breadth, 10.1; breadth
of brain-ease, 10.0; interorbital constriction, 3.7.
Distribution.—The type specimen was collected by F. Stephens
at Agua Caliente [= Palm Springs], Colorado Desert, Riverside
County, California, March 27, 1885. The only other Californian
A skull of the pocketed bat (U.S. Nation. Mus. ), from Fort
record of this bat, of which the writer is aware, is that by Elliot
(1904a, p. 321), who reports the finding by Edmund Heller of a
mummified specimen impaled on a mesquite bush in Palm Cajon,
near Palm Springs. The bush overhung a pool of water where bats
came to drink. (See map, text-fig. O.)
Natural History.—Nothing is known of the habits of the pocketed
bat.
Nyctinomus mexicanus (Saussure)
Mexican Free-tailed Bat
M{[olossus|. mexicanus Saussure (1860, pp. 283-285, pl. 15, figs. 2, 2a).
Original description; from Ameca, Jalisco, Mexico.
Nyctinomus nasutus, H. Allen (1864, pp. 7-10). Description; nomen-
clature; record of specimen from Fort Yuma.
Nyctinomus nasutus, Cooper (1868, p. 5). Distribution.
Nyctinomus nasutus, Cooper (in Cronise, 1868, p. 442), part. Occurrence
in California.
Nyctinomus nasutus, Coues and Yarrow (1875, pp. 81-82), part. Dis-
tribution.
Nyctinomus brasiliensis, Dobson (1878, pp. 437-439, pl. 32, fig. 8). De-
scription; general range.
Nyctinomus brasiliensis, True (1885, p. 603). Range.
Nyctinomus mohavensis, Bryant (1891a, p. 359). Distribution.
Nyctinomus mohavensis, Bryant (1892, p. 220). Nominal.
Nyctinomus brasiliensis, H. Allen (1894, pp. 163-171, pls. 32-33). De-
scription; nomenclature; distribution.
Nyctinomus brasiliensis californicus H. Allen (1894, p. 166, pl. 32, fig. 5).
Description of a ‘‘variety’’ from ‘‘California.’’ The only California
locality mentioned under N. brasiliensis is Fort Yuma.
Nyctinomus mohavensis, Stowell (1894, pp. 362-364). Deseription; dis-
tribution; habits of colony observed at San Jose, and occurrence at
Pacifie Grove.
Nyctinomus mohavensis, Elliot (1901, p. 418). Diagnosis.
Nyctinomus brasiliensis californicus, Miller and Rehn (1901, p. 272). Type
locality.
Nyctinomus mohavensis, Miller and Rehn (1901, p. 273). Type locality.
Nyctinomus mexicanus, Elliot (1904a, pp. 320-821). Records of oceur-
rence at Palm Springs and Fort Tejon.
Nyctinomus mexicanus, Elliot (1904b, pp. 629-630). Description; distri-
bution.
362 University of California Publications in Zoology (Vow. 17
Nyctinomus cynocephalus californicus, Stone (1904b, p. 587). Record of
occurrence at Belmont.
Nyctinomops mohavensis, Stephens (1906, pp. 273-274). Description;
distribution; habits.
Nyctinomus mexicanus, Elliot (1907, pp. 523-524). Record of specimens
taken at Fort Tejon, Palm Springs, and San Jose.
Nyctinomus mexicanus, J. Grinnell (1913b, p. 283). Range in California.
Nyctinomus mexicanus, J. Grinnell (1914, p. 268). Occurrence along
Colorado River.
Diagnosis —Size medium (total length 90 to 103 millimeters) ;
ears broad, apparently united at base, with a series of wart-like pro-
jections on anterior border; tragus small, flattened, squarely truncate
above; exserted portion of tail about equal to
that enclosed in membrane; general color, hair
brown.
Description: Head.—Ears about as broad as
long (text-fig. W), diverging from median line on
tance of 1 millimeter, although usually separate;
‘ ‘“‘warts’’ on anterior margin of ear varying in
view of head of .
REGRGRITO GEE number from five to eleven; a sinuous, sharply de-
icanus (drawn fined ridge arises from behind tragus and fades
from specimen no. away toward anterior external portion of lobe;
18921), X 1.00, hosterior surface of ear marked by five or six
showing rounded eens ‘ ~ 2
Reis, SA Se distinet transverse wrinkles; upper lip deeply
projections on an- crimped, forming perpendicular wrinkles. A deep
terior margin of furrow under eye.
Sls) SAE ace: Limbs and Membranes —Wings relatively small
ee Speed fe and weak (as compared with those of other Cali-
Arvil perpendicular fornia bats which equal this species in size of
wrinkles in upper body), slender; third metacarpal 1.7 length of
lip. fifth metacarpal. Wing attached to tibia on distal
third. Feet stout with toes distended.
Pelage.—Dorsal surface of body well furred with short soft hairs
averaging 3 or 4 millimeters in length. Hairs on ventral surface
slightly longer and more closely set. On face and chin, in addition
to shorter hairs, are scattered longer ones varying in length from 5
to 7 millimeters. Along edge of upper lip is a row of short, stiff,
downward-directed hairs. Ears very seantily furred with minute
hairs. Wings naked save for a narrow band of fur, continuous with
that of body, which extends from middle of humerus alone wing
to proximal third of femur, on both dorsal and ventral surfaces, and
for a fringe of hairs which extends along edge of post-calearial lobe
from its proximal end half way to its distal end. Outer edges of
first and fifth toes closely fringed with short, curved, white hairs; on
fifth toe, dorsal to this outer fringe, is a row of less numerous curved
hairs, three times as long as those of the first row. From upper
surface of base of each claw spring three or four long curved hairs
4 to 6 millimeters in length.
Color.—There is but little individual variation in color among the
series of Nyctinomus mericanus in the Museum of Vertebrate Zoology.
Fig. W. Side
forehead, and sometimes united at bases for a dis- —
1918] Grinnell: A Synopsis of the Bats of California 363
An example (no. 19006, 2) from Concord, Contra Costa County,
presents what is apparently the usual coloration of the species in
California. This specimen is hair brown in color, slightly paler
beneath, and with feet and membranes varying toward chaetura
black. The hairs on the body are everywhere whitish at their bases.
The long scattered hairs upon the face and chin are a dark hair
brown, while the long curved hairs upon the dorsal surface of the
claws are whitish.
Several specimens from Los Banos, Mereed County, and the two
examples from Painted Rock, Carrizo Plains, San Luis Obispo
County, exhibit a yellowish suffusion, due perhaps to absorption of
oil from the skin. Among thirty-one specimens from the vicinity
of Fresno, Fresno County, a single example shows marked departure
from the normal coloration. This individual is unusually sooty in
appearance, fur and membranes being chaetura black in color.
Skull—Brain-ease slightly rounded; sagittal crest very feebly
developed ; lachrymal and supraorbital ridges low but distinct; dorsal
surface of rostrum with a shallow longitudinal median coneavity ;
zygomata slightly expanded at middle. Dorsal profile nearly straight,
rising gradually from nose to top of brain-case; a depression between
top of brain-case and occipital ridge. (See pl. 23, fig. 48; pl. 24,
fig. 56).
= = = 9) OL
Teeth.—Dental formula: is ory s os, pm 99° m 54 = 30 or
32. Upper incisors simple, well developed, about half as high as
canines, wide apart at base, strongly converging at tips, separated
from canines by a space about equal to their greatest diameter, shaft
narrowing both above and below slightly developed cingulum, its
apex blunt. First and second lower incisors equal, their crowns in
contact with each other and with cingulum of canine. In young
specimens the crowns are bifid, with lobes rounded at the tip. In
old specimens the crowns are squarely truncated; in some all trace
of the notch is obliterated. Third incisor when present varies from
a mere spicule to a slender well-formed tooth, bifid at crown, and
three-fourths height of second incisor. In a series of thirty-seven
skulls belonging to the Museum of Vertebrate Zoology sixteen were
found to have three lower incisors on each side, eight to have two
only, four to have two in the right mandibular ramus, and three in
the left, and two to have three in the right ramus and two in the left.
Seven of the skulls were found to have less than four lower incisors,
the ones present being badly worn and broken.
Stowell (1894, p. 363) found that of forty-five specimens examined
twenty-four possessed six lower incisors, nine had five, and twelve
had four. He remarks:
The outer incisor when present is very small, and so crowded forward as to
occupy a precarious position in front of the canine, a fact which may account
for its absence in so many specimens. In by far the majority, the incisors are
distinctly bilobate and the lobes have well-rounded tips; but in some specimens
the tips have become more or less worn, and in a few individuals this process
has proceeded so far that the upper edges of the teeth are truncate, with scarcely
a trace of the median notch. In the specimens examined we have noticed that
most of those with perfectly truncate incisors have the latter also reduced in
number. This probably indicates that both conditions are dependent upon age.
364 University of California Publications in Zoology [Vou.17
While most of the skulls in the Museum of Vertebrate Zoology lack-
ing the third incisor, show badly worn teeth, yet two with unworn
teeth show no trace of the third incisor, indicating that the absence
of this tooth is sometimes a matter of individual variation, as well as
of age.
Canines strong and well developed, with distinct cingula. First
upper premolar minute; second well-developed. First lower premolar
about half as high as second, but with base nearly as large. Molars
normal,
Measurements.—Average and extreme measurements in milli-
meters of twenty examples of N. mexicanus in the Museum of Verte-
brate Zoology are as follows: ten males: total length, 95.4 (90.0-99.0) ;
tail vertebrae, 34.6 (32.0-88.0) ; tibia, 12.3 (11.6-13.0); foot, 10.1
(8.0-12.0) ; forearm, 41.4 (40.7-44.0) ; greatest length of cranium,
17.1 (16.6-17.5); zygomatic breadth, 10.0 (9.6-10.5); mastoid
breadth, 9.4 (9.3-9.7) ; interorbital constriction, 3.9 (3.9-4.2); ten
females: total length, 96.2 (91.0-103.0) ; tail vertebrae, 35.5 (31.0-
40.0); tibia, 12.2 (11.5-13.8); foot, 9.8 (8.0-11.0); forearm, 41.2
(39.0-43.0) ; greatest leneth of cranium, 16.9 (16.7-17.4) ; zygomatic
breath, 9.8 (9.6-10.0); mastoid breadth, 9.2 (9.1-9.5); interorbital
constriction, 3.9 (3.7-4.2).
Synonymy and History—Nyctinomus mexicanus was described
under the name Molossus mexicanus by Saussure (1860, p. 283) from
material obtained at Ameca, Jalisco, Mexico (Miller, 1912, p. 70).
Merriam (1889), p. 25) described a bat which appears to be identical
with N. mexricanus, from Fort Mohave, Mohave County, Arizona,
under the name Nyctinomus mohavensis. Harrison Allen (1894.
p. 166) deseribed the same species from California under the name
Nyctinomus brasiliensis californicus. The California bat, however,
appears to be distinet from brasiliensis of South America. The
writer has examined the five bats from Belmont, California, recorded
by Rehn (in Stone, 19046, p. 587) under the name Nyctinomus cyno-
cephalus californicus and finds them distinet from true cynocephalus
and in no way different from examples of mecicanus from elsewhere
in California.
Distribution.—N yctinomus mexicanus has been recorded from the
voleano Popoeatepetl, Mexico, northward through the United States
to Neweastle, Garfield County, Colorado (Warren, 1910, p. 286), and
eastward from the Pacifie to the middle of Texas where, according to
Bailey (1905, p. 215) ‘‘its eastern limit of range, so far as known,
agrees closely with the eastern limits of mesquite.’’ In California
the species is abundant throughout the Upper and Lower Sonoran
zones, from the southern border of the state northward at least to
Marysville Buttes, Sutter County. Throughout the central valleys
365
f the Bats of California
: A Synopsis o
Grinnell
1918]
Mus.
no.
3872
3875
6701
5135
5127
8317
14656
1465
6700
14655
12670
5129
5130
5133
5127
14665
22101
22100
14666
18501
MEASUREMENTS IN MILLIMETERS OF TWENTY EXAMPLES OF NYCTINOMUS MEXICANUS (SAUSSURE), FROM CALIFORNIA
Sex
+0 +0 +0 +0 +0 40 40 +0 40 100, 0, 2, OK QA DA OY OV AA OY
Locality
Palo Alto, Santa Clara Co.
Palo Alto, Santa Clara Co.
San Francisquito Canon, Los Angeles Co,
Santa Monica, Los Angeles Co.
Pasadena, Los Angeles Co.
Sweetwater Dam, San Diego Co.
Los Banos, Merced Co.
Los Banos, Merced Co.
Glendora, Los Angeles Co.
Los Banos, Merced Co.
Fresno, Fresno Co.
Pasadena, Los Angeles Co.
Pasadena, Los Angeles Co.
Pasadena, Los Angeles Co.
Pasadena, Los Angeles Co.
Carrizo Plains, San Luis Obispo Co.
Coulterville, Mariposa Co.
Coulterville, Mariposa Co.
Carrizo Plains, San Luis Obispo Co.
Marysville Buttes, Sutter Co.
vertebrae
Tail
Foot
11.0
11.0
8.0
8.0
8.0
11.0
12.0
11.0
9.0
12,
9.5
8.0
9.0
8.0
8.0
11.0
9.0
9.0
11.0
11.0
Forearm
41.0
41.0
41.0
41.3
44.0
41.0
40.7
41.0
41.3
41.9
40.5
39.0
40.0
41.4
42.5
43.0
42.9
o.4
41.5
40.0
42.9
2.8
» Greatest
length of
cranium
=
Zygomatic
breadth
Mastoid
breadth
constriction
co wo oo x & Interorbital
C=)
ie
°
366 University of California Publications in Zoology (Vou. 17
of the state it is one of the most abundant bats, occurring in great
colonies where conditions are favorable. (See map, text-fig. X.)
Specimens Examined.—The writer has examined 217 specimens
of Nyctinomus mexicanus from the following localities in California:
@ NYCTINOMUS MEXICANUS
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY
! UNIVERSITY OP CALIFORNIA
Fig. X. Map showing stations of occurrence in California of Nyctinomus
mezxicanus, as established by specimens examined by the author.
San Diego County: Sweetwater Dam, 1; Imperial County: 8 miles
east of Picacho, 1; Echo Island, Salton Sea, 1; Riverside County:
Mecea, 1; San Jacinto, 10 (Stanford Univ.) ; San Bernardino County:
Chemehuevis Valley, 1; San Bernardino, 2; Los Angeles County:
Glendora, 1; Pasadena, 11; San Francisquito Canon, 1; Santa Moniea,
1918] Grinnell: A Synopsis of the Bats of California 367
1; Ventura County: Santa Paula, 1 (Stanford Univ.) ; Ventura, 8;
Kern County: Fort Tejon, 1; San Luis Obispo County: Painted Rock,
Carrizo Plains, 2; Fresno County: Fresno, 29; Lane Bridge, 10 miles
“north Fresno, 2; Mariposa County: El Portal, 1; Coulterville, 2;
Mono County: Walker Lake, 1; Merced County: Los Banos, 18;
Stanislaus County: Grayson, 1; Santa Clara County: Menlo Park, 7;
Palo Alto, 13 (Stanford Univ., 5; Mus. Vert. Zool., 8); Stanford
University, 1 (Calif. Acad.); San Jose, 52 (Stanford Univ.); San
Mateo County: Belmont, 5 (Acad. Nat. Sei. Phila.) ; Alameda County :
Berkeley, 2 (U. C. Dept. Zool.) ; Alameda, 1; San Joaquin County:
Tracy Lake, 6 miles southwest Galt, 1; Contra Costa County: Con-
cord, 24; Eldorado County: Limekiln, 12; Placer County: Auburn,
1; Sutter County: Marysville Buttes, 3 miles southwest Sutter, 1.
Natural History—This bat is gregarious and apparently prefers
retreats in buildings rather than those afforded by nature. I know
of no instance of the finding of these bats in natural haunts save
where they had no choice of other quarters. On May 27, 1911,
Swarth (MS) found a cluster of these bats in company with a large
number of Antrozous pacificus in a erevice extending upward from
a hollow at the base of Painted Rock, Carrizo Plains, San Luis Obispo
County. When a buggy whip was thrust up among them some of
the bats crawled farther up into the crevice, while about fifteen or
twenty tumbled out and flew about the rock seeking other shelter.
At Glendora, Los Angeles County, a raid upon a colony of Antrozous
occupying the spaces under the eaves of a barn loft resulted in the
capture of ten Antrozous and a single male Nyctinomus mexicanus.
More of the latter species may have been present as many of the bats
disturbed eluded capture and escaped into the open air. Stowell
(1894, pp. 362-364) records the finding of large numbers of these
bats behind the iron window shutters of the courthouse in San Jose.
On February 27, the leaves of one of these shutters were opened and
the bats were found thickly clustered in the darker recesses. The
bats made little effort to escape and about seventy specimens were
secured. These consisted of males and females in about equal num-
bers. The window from which these bats were secured was on the
west side of the building. On March 3, of the same year, thirty-two
specimens were taken from another west window and these bats also
consisted of an equal proportion of the two sexes. An examination
of four different windows on the east side of the building resulted in
the capture of thirty-five specimens, all females. W. P. Taylor
368 University of California Publications in Zoology (Vou. 17
P
(MS), when encamped in April, 1912, near Marysville Buttes, Sutter
County, noted bats of this species flying down the cafion each even-
ing. They appeared shortly after 6:45 p.w. OC. H. Richardson (MS)
took six male N. mexicanus from spaces between the sheathing and
shakes on a barn beside a slough near Los Banos, Merced County,
on March 25, 1911. He writes: ‘‘This was evidently an old roost,
for the dung lay thick upon the floor. There were many bats present,
to judge from the amount of squeaking and the odor.”’
Several specimens of this bat have been sent to the Museum of
Vertebrate Zoology from the attic of a schoolhouse at Fresno.
Heller (in Elhot, 1904a, p. 320) found Nyctinomus mexicanus
common in the spring of 1902 in Palm Canon, near Palm Springs,
Riverside County. He also found several hundred living in the
garret of one of the old buildings at Fort Tejon, Kern County.
Specimens secured during November and December from the attic
of a blacksmith shop at Concord, Contra Costa County, and sent to
the Museum of Vertebrate Zoology, were excessively fat. The males
examined had very small testes. Specimens secured from the same
colony January 31 were very thin. In one of the males examined the
testes were shghtly enlarged, indicating the possibility of spring
mating in this species. The stomachs of four specimens were
crammed with the remains of insects, flies and small beetles, too finely
triturated for identification. I do not think that these bats had
hibernated but that the loss of fat was due to the seareity of food,
caused by the frosty weather of December and January.
Along the Colorado River between Needles and Yuma, in the
spring of 1910, Museum collectors found the species not uncommon.
In the neighborhood of Mecca, Riverside County, the species was
found during the spring of 1908, and one specimen was collected from
a colony found in a cave on Echo Island in Salton Sea.
On the evening of January 1, 1915, J. Grinnell (MS) noted several
bats flying at El Portal, Mariposa County, 2000 feet altitude. One
of the two examples shot proved to be Nyctinomus mexicanus. This
specimen, a female, weighed 11.5 grams. Specimens and records at
hand establish the presence of the Mexican free-tailed bat in the same
general areas of the state throughout the year. It is therefore safe
to infer that this species is but slightly or not at all migratory.
On October 27, 1916, J. R. Pemberton and J. Grinnell investigated
a colony of bats (pl. 14, fig. 2) oceupying the attie of a brick house
at San Bernardino, the home of R. B. Herron. Dr. Grinnell records
(MS) his observations as follows:
1918 | Grinnell: A Synopsis of the Bats of California 369
I found just one species, Nyctinomus mexicanus, though Mr. Herron declares
there are at least three kinds present, at different seasons of the year. I went
all through the attic and all I saw distinctly, flying or in different ‘‘colonies,’’
were of the above species. There are now at least 1000 bats, clinging together
in patches in the peak of the roof, or crowded into crevices between the ridgepole
and the shingles. Along the floor is a longitudinal heap of guano 6 inches to
1% feet high. Herron says that he has taken out five grain sacks of the
guano at one time, and used it about the garden as fertilizer. The bats ‘‘chuckle’’
continually, a subdued chorus of talking sounds. There are very few of the
shrill squeaks. The animals crawl, back upwards, as far as they can get. The
lower ones crowd against the upper ones, the latter squeezing farther into the
crevices, the whole mass in continual agitation. When thoroughly disturbed
many took flight, and the fluttering wings and occasional collisions with me (or
attempts to alight) gave a weird impression.
Mr. Herron thinks these bats migrate—they seem to be entirely gone for three
or four months, in cold weather, and then all at once they come in. ‘‘In summer
time there are three kinds, a big broad-winged one, a little narrow-winged one,
and this one.’’
Sun sets at 4:57. Perfectly clear weather. First bat out at 5.04. At 5:07,
27 had come out. At 5:10, 95 more had come out. At 5:15, 120 more had
come out. At 5:30, when it had gotten too dark for me to count longer, 1367
more had come out. Total counted, 1609.
Although there were three gable ends, each with a small slatted ventilator
window, all the bats came out of the west-facing one. Mr. Herron says the
first usually comes out just before the sun sets, and all are out in about half an
hour. He thinks there are as many as 5000 there in midsummer. Just before
they came out, in little squads of 10 to 50, I heard a low chuckling chorus at
the grating; then the crowd began to pour out.
Two female Mexican free-tailed bats taken by Ferris and Storer
7
at Coulterville, Mariposa County, June 7, 1915, each contained one
large embryo.
Twelve specimens of Nyctinomus mexicanus secured by Mr. Joseph
Dixon at Limekiln, Eldorado County, between August 3 and 6, 1916,
were females and varied in weight from 11 to 15 grams. Dixon states
(MS) that the mammary glands of these bats were full of milk,
although the glands of adult individuals of Eptesicus and Myotis
taken at the same time were undeveloped.
Genus Eumops Miller
Ears very large, rounded or squarish in outline, joined across
forehead, usually extending shghtly beyond nostril when laid forward,
the anterior margin without horny processes; antitragus distinet but
not thickened, keel greatly developed, flattened and expanded at edge ;
tragus small, flat, its upper edge truncate or rounded. Lips full and
expanded, not conspicuously wrinkled. Muzzle pad well developed,
deeply emarginate above, its upper edge and median ridge with
minute horny processes and small spoon hairs. Wings, feet and tail
as given for the family.
370 University of California Publications in Zoology {|Vou.17
Leneth of skull shehtly more than twice its greatest width; ros-
trum more than half length of brain-case; angle between rostrum and
brain-case very slight, in fact whole dorsal profile nearly straight;
sagittal crest but slightly developed; zygomata not expanded at
middle; palate slightly arched.
iss = 9-9 = =
Dental formula: is, e =. pm 529 or = m = — 30 ore2or
Upper incisors long and slender, nearly one-half height of upper
canines and diverging slightly near their tips. Lower incisors tall,
slender, and crowded in a semicircular row beneath projecting cingula
of canines; crowns of lower incisors bifid and one-third wider than
columnar shafts. Canines large and robust (see pl. 20); height of
upper canine about one-third more than antero-posterior diameter at
cingulum; anterior face slightly grooved, and inner face somewhat
coneave ; lower canines slightly smaller than upper, but similarly pro-
portioned. First upper premolar, if present at all, minute and closely
crowded between canine and second premolar. Posterior upper pre-
molar with crown area more than half that of first molar and exceed-
ing it in height. Lower premolars with crowns about as long as
broad; that of second slightly exceeding that of first in height.
Molars not peculiar in form; crowns of lower exceed those of upper
in height. In Nyctinomus the upper incisors are wide apart at their
bases, with converging tips, whereas in Hwmops the upper incisors
are in contact at their bases, diverging toward the tips.
One species of Humops has been recorded from California.
3
Eumops californicus (Merriam)
California Mastiff Bat
Molossus californicus Merriam (1890, pp. 31-32). Original description;
type locality, Alhambra, Los Angeles County, California.
Molossus californicus, Bryant (1891a, p. 359). Nominal.
Molossus californicus, J. A. Allen (1891a, p. 198). Distribution; relation-
ships.
Promops perotis californicus, H. Allen (1894, pp. 175-182, pls. 36-38).
Description.
Promops californicus, Elliot (1901, pp. 422-423, fig. 94). Description;
distribution.
Promops californicus, Miller and Rehn (1901, p. 271). Type locality.
Promops californicus, Elliot (1905, p. 498). Geographic distribution.
Promops californicus, Stephens (1906, p. 275). Description; distribution.
Eumops californicus, Miller (1906, p. 85). Description (as type species
of new genus).
Eumops californicus, Miller (1912, p. 71). Type locality.
Eumops californicus, J. Grinnell (1913b, p. 283). Range in California.
Diagnosis.—Size large (total length 157 to 184 millimeters) ; ears
very broad, united above nostrils, directed forwards rather than
upwards; tail exserted far beyond edge of interfemoral membrane ;
upper incisors in contact at their bases, diverging toward tips. (See
pl. 19, figs. 19-20; pl. 20, figs. 21-23.)
1918 | Grinnell: A Synopsis of the Bats of California 371
Description—Head elongate (length of head, in alcoholic speci-
men, 42 millimeters, greatest width, 20); muzzle pad prominent,
deeply emarginated above, its upper edge and median ridge with
minute horny processes and small spoon-hairs; nostrils large, directed
downward and outward; eyes placed far back on sides of head (center
of eye, in alcoholic specimen, 21 millimeters from tip of muzzle), and
almost concealed by the drooping anterior border of the auricle. In
front of each eye is a prominent glandular swelling. Ears large,
directed outward and forward, joined in median line, and so large as
to entirely conceal face and muzzle from an observer when viewing the
bat from the dorsal side. Ear conch, or auricle, broadly convex anter-
iorly and posteriorly, slightly convex on top (pl. 20, fig. 21). Tragus
small, truncate (in aleohohe specimen 3.5 millimeters in height at
anterior edge and 3 millimeters in breadth at top), and nearly con-
cealed by the semi-oval antitragus, which is (in alcoholic specimen)
10.5 millimeters in length and 7 in greatest height; antitragus sep-
arated posteriorly from conch by a deep notch. Ears haired along
margin, but greater portion of keel, both externally and internally,
naked. Folds of ears directly above nostrils heavily haired on both
upper and lower surfaces. Upper lp with a thick fringe of down-
ward-direeted hairs.
Limbs and Membranes.—Wings long and slender (pl. 19); fifth
metacarpal scarcely more than half length of third. Dorsal surface
of wing membrane, as well as interfemoral membrane, furred to a
distance of about 12 millimeters from body. A small patch of hairs
on dorsal surface of antebrachial membrane, this patch extending
inward from edge of radius, except at bottom of angle formed by
junction of radius with humerus; another group of hairs extending
along distal three-fourths of posterior edge of forearm, widening and
occupying angle between forearm and fourth metacarpal; a third
small group of hairs at distal end of third metacarpal. A small patch
of fur on ventral surface of antebrachial membrane, occupying angle
between anterior edge of membrane and proximal half of humerus;
a line of fur extends from this patch across humerus and along its
posterior edge and unites with a small patch on lateral. membrane
immediately posterior to distal end of humerus. Interfemoral mem-
brane not haired, except for a narrow border along proximal margin
of its dorsal surface, and a few scattering hairs along edge of post-
ealearial lobe. Legs and feet short and stout; feet only twice as
long as wide.
Pelage—F ur long and soft, denser and shorter on abdomen than
elsewhere, varying in length from 5 to 10 millimeters.
Color—Membranes deep hair brown; fur a lhghter tone of same
color, slightly paler on ventral surface; bases of hairs everywhere
whitish for a distance of from one-half to three-fourths their length.
Skull and Teeth.—As given for the genus (see p. 370).
Measurements.—Average and extreme measurements in milli-
meters of ten specimens of the mastiff bat in the Museum of
Vertebrate Zoology are as follows: total length, 167.1 (extremes,
157.0-184.0) ; tail vertebrae, 59.7 (52.0-70.0) ; tibia, 22.1 (21.0-23.3) ;
foot, 16.8 (13.0-19.0) ; forearm, 72.0 (69.0-74.5) ; greatest length of
cranium, 31.0 (30.0-32.0). Five specimens, measured fresh, showed
3712 University of California Publications in Zoology (Vou. 17
stretch of wings, tip-to-tip, of 526 millimeters (515-535) [== 20-21
inches].
MEASUREMENTS IN MILLIMETERS OF TEN EXAMPLES OF EUMOPS CALIFORNICUS
(MERRIAM), FROM CALIFORNIA
Fe 2 é Fea RIE
no. Sex i= a =) i is N a)
43266 J 165.0 (GSO eee 15.0 73.5 32.0 17.8 15.3
5242? 157.0 55.0 22.0 18.0 69.4 31.9 I(es3) 14.8
16597 184.0 70.0 23.0 17.0 W200. 2h ae eS
43277 9 157.0 Gi ee 14.0 71.0 31.2 17.6 14.7
6946" 9° 167.0 55.0 23.3 13.0 69.0 30.0 17.0 14.8
19021° 9 171.0 59.0 21.0 19.0 71.0 30.8 17.4 15.1
190227 9 166.0 52.0 23.0 17.0 74.5 30.6 17.0 14.6
191917 Q 173.0 63. 21.1 18.0 72.0 31.0 18.0 15.1
19192? 9° 166.0 61.5 21.4 18.5 73.4 30.0 18.2 15.3 =
192837 9 165.0 ORO! | seer 19.0 74.5 Seri 18.0 15.2 5.2
1From Pasadena, Los Angeles County.
?From Sierra Madre, Los Angeles County.
* From Colton, San Bernardino County.
Synonymy and History—This bat was described by Merriam
(1890, pp. 31-32) from a specimen taken at Alhambra, Los Angeles
County, California, under the name Molossus californicus. Miller
(1906, p. 85) made Merriam’s Molossus californicus the type species
of the new genus Humops, distinguished from the nearly related
genera Molossus and Promops by reason of its arched rather than
domed palate and less well-developed sagittal crest.
Distribution.—Miller (1907, p. 256) figures the skull of a specimen
taken at Tucson, Arizona, and Bailey (1905, p. 216) records the
capture of a specimen of LZ. californicus at the bottom of Pump
Cafion, near Langtry, Texas. J. Grinnell (1913b, p. 284) gives the
range of this bat in California as follows: ‘‘Lower Sonoran zone of
southern California; most numerous in the San Diegan district, but
noted also on the Colorado Desert, and in the San Joaquin Valley,
in Kern and Fresno counties (Mus. Vert. Zool.) ; northernmost station,
Fresno.’’ The basis of the latter record was two specimens seen in
a taxidermist’s shop in Fresno, and stated to have been caught in
the courtyard of that city. (See map, text-fig. H.)
Specimens Examined.—tThe writer has examined 26 examples from
the following localities in California: San Diego County : Dos Cabasas,
1 (U.S. Nation. Mus.) ; Otay, 1 (Stanford Univ.) ; Riverside County:
Mecea, 1; San Bernardino County: Colton, 1; Los Angeles County:
1918 ] Grinnell: A Synopsis of the Bats of California 373
Los Angeles, 2 (U.S. Biol. Surv., 1; U. S. Nation. Mus., 1) ; Pasadena,
6; Sierra Madre, 9; Kern County: Sumner, 1; Buttonwillow, 1
(Calif. Acad. Sci.) ; Bakersfield, 1 (Univ. Calif. Dept. Zool.) ; Tulare
County: Traver, 1 (Stanford Univ.) ; Fresno County: Fresno, 1.
Natural History.—The type specimen of EL. californicus was found
by E. C. Thurber on a December evening on a ledge over a door.
Merriam (1890, p. 31) says: ‘‘Two others were caught during the
same month (December, 1889), and both in similar situations. Mr.
Thurber says of one of them: ‘It was hanging from the ledge of a
window, swinging back and forth and knocking against the window
as if to attract attention. All were caught about 8 or 9 o’clock in
the evening.’
On October 1, 1916, Adrey Borell, of Fresno, sent to the Museum
of Vertebrate Zoology a live mastiff bat which he had just found
hanging on the inside of a window-sill in a school building in Fresno.
H. C. Ohl (MS) found a mastiff bat in a railroad round-house at
Mendota, Fresno County, in December, 1911. The last named locality
constitutes the northwesternmost record station to date.
Stephens (1906, p. 275) records the capture of a specimen behind
a sign board and another in a tunnel, and adds that all dates known
to him are in winter.
On or about October 1, 1907, a specimen of EF. californicus (now
no. 23391, Mus. Vert. Zool.) was found clinging to the side of a house
at Mecea, Riverside County.
On March 8, 1909, W. B. Donnell discovered five examples of
E. californicus in Pasadena in the attie of an old house which was
being torn down. These bats were very much emaciated, and the
stomachs were found to be empty. These specimens are now nos.
4326-4328, and 94388-9439 (Mus. Vert. Zool.).
On December 27, 1912, at the writer’s suggestion, Mr. Charles
L. Camp, of Sierra Madre, Los Angeles County, kindly investigated
a rookery where he had found bats common in the summer time. This
haunt was an old shedlike structure, some sixty feet long, and three
stories high. The interior of the building was very dark and con-
tained piles of old fruit-drying trays. In the summer time some bats
had been found in between the trays, but more were discovered hang-
ing to the shingles and rafters in the east gable of the building. The
December visit revealed but a single living bat in the building, an £.
californicus. This specimen was found hanging from the rafters.
Near it hung a dead bat of the same species, evidently just killed and
374 University of California Publications in Zoology {[Vou.17
perhaps a victim of the unusually cold night which preceded the eol-
lector’s visit. The mummified remains of a third individual of the
species was found upon a heap of guano upon the floor of this gable,
together with the dried remains of two specimens of Antrozous
pacificus. A visit to the same place made on January 6, 1913, failed
to reveal any bats.
The live Humops californicus secured by Mr. Camp, December 27,
1912, at Sierra Madre was received at the Museum of Vertebrate
Zoology, December 28. She hung herself up by the feet in a dark cor-
ner of the cage in whieh she was domiciled, and remained quiet until
forcibly removed. She used feet and half-closed wings in an energetic
endeavor to dislodge the writer’s fingers from her back, and constantly
kept her mouth opened to the widest extent in an effort to seize the
fingers, all the while uttering shrill cries, loud enough to be heard at a
distance of several hundred feet. These sounds might be likened to
the utterances of a young robin in distress, and were cries, not mere
squeaks.
When placed in a glass jar containing a bit of cotton saturated
with ether the bat made strenuous efforts to escape, striking out with
both feet and half-closed wings. While the bat was still partially
under the influence of the anesthetic an endeavor was made to smooth
her dampened and ruffled fur with a small brush. She curled and
twisted about the offending object in a way which showed the greatest
flexibility of body. When left to her own devices the bat clung
- to the rod above her with her right foot while with her left foot she
thoroughly combed the dorsal surface of her left wing and her head
and body on the left side, both dorsally and ventrally. Then shift-
ing her hold to the other foot she repeated the combing process for
the right half of her head and body with the alternate foot. Finally
she became quiet and hung for a time motionless, her whole body
being supported by one foot, the other hanging down on her back,
the claws toward the center of her back and the folded ankle joint
resting in a pocket formed by a fold of the wing membrane. Her
eyes remained open but were completely in the shadows cast by the
anterior margins of her ears and her loosely folded wings.
When the bat was placed gently upon the floor of her cage she
scuttled backward with surprising rapidity, backing up the side of
the cage until her nose just cleared the floor. She remained for
some time in this position supporting herself by clinging with her
claws to the unplaned surface of the board. When the observer
1918] Grinnell: A Synopsis of the Bats of California 375
returned to the cage an hour later the bat was found hanging head
downward in a dark corner. In this position she remained for several
days, and seemed absolutely indifferent to particles of fresh meat or
insects held very near to her mouth.
When placed upon a smooth rug the bat scrambled forwards using
her folded wings with great dexterity. In this instance the terminal
joints of the wing were tucked away close to the bat’s body, beneath
the lateral membrane. When placed upon pebbly and twig-strewn
ground the bat, before starting forwards, folded her slender wings
closely and placed the delicate terminal portion upon the dorsal sur-
face of the wing itself, in the angle between the humerus and radius,
safe out of harm’s way. After being kept alive in captivity for
three weeks the bat was chloroformed, as it refused to eat. The
weight of the living bat when first received was found to be 114 ounces
(42.5 grams).
In the latter part of May, 1913, Mr. Camp procured three more
living Humops from the same Sierra Madre haunt. These were kept
alive at the Museum of Vertebrate Zoology from May 24 to June 5,
1913, and were then chloroformed, as they unfailingly ignored all food
offered them. Hach individual proved to be a female containing a
single embryo. The breeding time of the species is thus indicated.
376 University of California Publications in Zoology (Vou. 17
LITERATURE CITED
ACKERT, J. E.
1914. The innervation of the integument of Chiroptera. Jour. Morph., 25,
301-343, 4 pls.
ALLEN, G. M.
1916. Bats of the genus Corynorhinus. Bull. Mus. Comp. Zool., 60, 333-356,
7 figs. on 1 pl.
ALLEN, H.
1862. Descriptions of two new species of Vespertilionidae, and some remarks
on the genus Antrozous. Proc. Acad. Nat. Sci. Phila., 1862, 246-
248.
1864. Monograph of the bats of North America. Smithsonian Mise. Coll.,
7, pp. 1-xxiii, 1-85, 68 figs. in text.
1866. Notes on the Vespertilionidae of tropical America. Proc. Acad. Nat.
Sci. Phila., 1866, 279-288.
1891. On a new species of Atalapha. Proc. Amer. Philos. Soe., 29, 5-7.
1892. A new genus of Vespertilionidae. Proc. Acad. Nat. Sci. Phila., 1881
[=Jan. 19, 1892], 467-470.
1894. A monograph of the bats of North America. Bull. U. 8. Nat. Mus.,
43 (1893), pp. ix+198, 38 pls., 1 fig. in text.
ALLEN, J. A.
1890. Notes on collections of mammals made in central and southern Mexico,
by Dr. Audley C. Buller, with descriptions of new species of the
genera Vespertilio, Sciwrus, and Lepus. Bull. Amer. Mus. Nat.
Hist., 3, 175-194.
1891a. Description of a new species of big-eared bat, of the genus Histiotus,
from southern California. Bull. Amer. Mus. Nat. Hist., 3, 195-198.
1891b. On a collection of mammals from southern Texas and northeastern
Mexico. Bull. Amer. Mus. Nat. Hist., 3, 219-228.
1893a. List of mammals collected by Mr. Charles P. Rowley in the San Juan
region of Colorado, New Mexico and Utah, with descriptions of new
species. Bull. Amer. Mus. Nat. Hist., 5, 69-84.
1893b. On a collection of mammals from the San Pedro Martir region of
Lower California, with notes on other species particularly of the
genus Sitomys. Bull. Amer. Mus. Nat. Hist., 5, 181-202.
1894. Deseriptions of ten new North American mammals, and remarks on
others. Bull. Amer. Mus. Nat. Hist., 6, 317-332.
1895. On a collection of mammals from Arizona and Mexico, made by
Mr. W. W. Price, with field notes by the collector. Bull. Amer.
Mus. Nat. Hist., 7, 193-258, 17 figs. in text.
1906. Mammals from the states of Sinaloa and Jalisco, Mexico, collected
by J. H. Batty during 1904 and 1905. Bull. Amer. Mus. Nat. Hist.,
22, 191-262, pls. 20-33.
Autom, B.
1863. Die Nahrung unsere Eulen. Journal fiir Ornithologie, 11, 41-46, 217—
219.
ANDERSEN, K.
1912. Catalogue of the Chiroptera in the collection of the British Museum.
Ed. 2, 1, pp. ci+ 854, 79 figs. in text.
1918] Grinnell: A Synopsis of the Bats of California 377
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1918 | Grinnell: A Synopsis of the Bats of California 381
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382 University of California Publications in Zoology {|Vou-17
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Transmitted March 22, 1917.
eo
, a
PLATE 14
Fig. 1. Portion of Colorado Desert near Toro, Riverside County, showing
entrance to cave occupied by colony of California leaf-nosed bats (Macrotus
californicus). This eave and other similar ones in the vicinity also inhabited
by bats were formed by wave action at the margin of the ancient Blake
Sea. From these day-time retreats the bats issue forth at late dusk for
nocturnal foraging over the adjacent desert.
Fig. 2. View in attic at home of R. B. Herron, Colton, San Bernardino
County, showing clusters of Mexican free-tailed bats (Nyctinomus mexicanus)
crowded together under the ridgepole in the peak of the roof. At the time
of the observer’s visit these bats were continually crowding and _ jostling
each other in an apparent effort to squeeze still farther into the crevices.
Other bunches of bats of the same species were seen in different parts of the
garret, and at evening as the bats left this retreat through a slatted ventilator
at the gable end, 1609 were counted. The photograph here reproduced was
taken on October 27, 1916. Mr. Herron states that in midsummer the colony
is much larger, including, he thinks, about 5000 individuals. Flashlight
photograph by J. R. Pemberton.
Fig. 3. Station in the Transition life-zone near Fyffe, Eldorado County,
with an incense cedar in foreground and yellow pines in background. Species
of bats collected here July 19-31, 1916, were: Lasionycteris noctivagans, Myotis
californicus californicus, Myotis longicrus longicrus, Myotis lucifugus altipetens,
Myotis thysanodes, Eptesicus fuscus, and Pipistrellus hesperus merriami. This
photograph shows an ideal collecting site, a tongue of cleared land projecting into
a forested area.
[ 384 ]
UNIV. CALIF. PUBL. ZOOL. VOL. 17
PLATE 15
Figs. +, 5, 6. Photographs of a living lump-nosed bat (Corynorhinus rafi-
nesqua pallescens, no. 21435, g), secured in a cave near Whitewater, Riverside
County. 1.00. Fig. 4 shows the bat in a dormant state, with wings held
close to the body and ears folded back under them. The pinnae here are
not bent sharply back, but curved by a regular and even crimping of the
posterior edges of the auricles, the resulting appearance reminding the observer
of the curving horns of a mountain sheep. Fig. 5 was taken when the animal
had been disturbed and seemed to be endeavoring to overcome the lethargy
of profound slumber. Fig. 6 shows the bat fully aroused and about to take
flight. The successive positions of the ears illustrate the way in which they
are unfurled.
[ 386 |
UNIV. CALIF. PUBL. ZOOL. VOL. 17 (GRINNELL] PLATE |
Fig. 4
Fig. 6
er
PLATE 16
Fig. 7. Hoary bat (Nycteris cinerea, no. 21439, 9), secured at Fresno.
Photographed under chloroform. Ventral view, X 0.43, showing tail curved
over body ventrally. Note the furring of the dorsal surface of the inter-
femoral membrane and of the anterior portion of the ventral surface of the
wing membranes.
Fig. 8. Hoary bat (Nycteris cinerea, no. 20778, J) front view, X 1.20. This
figure shows the form of the ear and tragus, and the distribution of hair
on the ventral surface of the ear; also the tumid-rimmed and widely separated
nostrils.
Fig. 9. Spotted bat (Huderma maculatum, no. 1196), found dead in overflow
from railway water tank, at Mecca, Riverside County. Photograph from dried
skin. Dorsal view, X 0.41. Note the large ears, the parchment-like wing and
tail membranes devoid of fur, and the striking color-pattern of the back. This
coloration reminds the observer of the ‘‘death’s head’’ pattern displayed upon
the thorax in certain moths.
[ 388 ]
UNIV. CALIF. PUBL. ZOOL
PLATE 17
Fig. 10. Hoary bat (Nycteris cinerea, no. 20778, g), photographed under
chloroform. Dorsal view, X 0.90. Note the large stout feet and thumbs; the
thickly furred back and interfemoral membrane; the three small patches of
light-colored fur, one on the antebrachial membrane at the proximal fifth of
the forearm, one at the base of the fifth metacarpal, and one at the base of
the thumb; and the long narrow wings, the third metacarpal far exceeding
the fifth in length. The unusual length of the tail in this genus is not shown,
as in this photograph the tip is turned under the body (see pl. 16, fig. 7).
Fig. 11. Pacific pallid bat (Antrozous pacificus, no. 9440, 3), secured at
Fresno. Photographed under chloroform. Dorsal view, X 0.50. The ears
of this bat are separate and wide apart at base, and extend considerably
beyond the tip of the snout. The feet are stout and more than half the length
of the relatively short tibiae. The wings are short and broad, the third
metacarpal being but slightly longer than the fifth.
Fig. 12. Lump-nosed bat (Corynorhinus rafinesquit pallescens), X 0.87. The
bat is asleep, hanging head downward, and is supported mainly by the claws,
which are thrust into a crevice in the angle where two boards join. The
thumbs are turned back under the wings, and their claws thus aid in support-
ing the weight of the animal. One of the tragi shows clearly, as do the two
glandular swellings on the snout.
[ 390 ]
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PLATE 18
Fig. 13. Large brown bat (Hptesicus fuscus, no. 22084, ¢), X 0.62. Dorsal
view of fresh specimen secured in Yosemite Valley, Mariposa County. Note
the heavily-built forearm, tibia and foot, and the stout thumb. The bluntly
rounded ears of this bat are somewhat longer than broad, a fact not apparent
in the photograph.
Fig. 14. Red bat (Nycteris borealis teliotis, no. 21516, 2), XK 0.55. Dorsal
view of fresh specimen secured at Fresno. Note the small foot and ear, and
the long interfermoral membrane, haired over the entire surface. The char-
acter of the dorsal pelage is well shown in this photograph: the frost-like light
tippings of the hairs, and the short dense fur on the head as compared with
the longer hair on the back.
Fig. 15. High Sierra bat (Myotis lucifugus altipetens, no. 23035, ¢), X 0.73.
Dorsal view of fresh specimen taken at Vogelsang Lake, Mariposa County.
Note the form of the ear (longer than broad and bluntly rounded at tip),
the long slender thumb and the relatively long foot.
Fig. 16. Dorsal view of fresh specimen of Merriam bat (Pipistrellus hesperus
merriami, no. 22081, ¢), secured at Pleasant Valley, Mariposa County; X 0.81.
This bat and the one shown in fig. 17, Myotis c. californicus, are of somewhat
similar size, but it will be noted that the ears of californicus are of nearly the
same tone of coloration as the back, whereas the ears of merriami are of a
blackish shade, contrasting with the grayish tone of the back. The ear in mer-
riami is seen to be shorter than in californicus.
Fig. 17. Dorsal view of fresh specimen of little California bat (Myotis cali-
fornicus californicus, no. 22078, g), secured at Pleasant Valley, Mariposa County.
X 0.81.
Fig. 18. Living adult female red bat (Nycteris borealis teliotis, no. 24179)
with her three young; X 0.44. This bat was found at Stockton, San Joaquin
County, July 12, 1915, clinging to the trunk of a tree one and one-half feet above
the ground, with the young all attached to her. Her weight was found to be
12.4 grams and the combined weights of the three young, 19.4 grams.
UNIV. CALIF, PUBL. ZOOL. VOL. 17 [GRINNELL] PLATE 18
PLATE 19
Fig. 19. Mastiff bat (Humops californicus, no. 19022 [or 1], 2); X 0.41.
Dorsal view of individual taken at Sierra Madre, Los Angeles County. Photo-
graphed under chloroform. Note the large ears, joined at their anterior bases;
the long narrow wings, the fifth metacarpal being scarcely more than half the
length of the third; the greatly exserted tail; the different quality of the pelage
on the dorsal surface of the head from that on the back; the stout foot; and the
short tibia.
Fig. 20. Mastiff bat (Huwmops californicus, no. 19283 [or 2], 2); X 0.58.
Ventral view of individual taken at Sierra Madre, Los Angeles County. Photo-
graphed under chloroform. Note the downward-directed nostrils, huge mouth,
wrinkled ears, long, slender wings, and stout thumbs.
[ 394 ]
UNIV. CALIF. PUBL. Z
3L. ZOOL. VOL. 17
[GR NNELL ]
Fig. 20
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PLATE 20
Fig. 21. Mastiff bat (Hwmops californicus, no. 19021 [or 2], 2); X 1.52.
View of head with mouth opened to show dentition. Photograph of fresh speci-
men secured at Sierra Madre, Los Angeles County. Note the large ear directed
outward and forward, the small truncate tragus, and the semi-oval antitragus
which is separated from the rest of the auricle by a deep notch. The form and
position of the nostrils is here well shown. It will be seen that the two upper
incisors are long, slender, and in contact at their bases, though diverging at
the tips.
Fig. 22. Dorsal view of skull of Huwmops californicus, no. 4327, 2. X 2.
Fig. 23. Side view of skull of Hwmops californicus, no. 4327, 9. X 2.
[ 396 ]
PLATE 21
Dorsal views of the skulls of twelve species of California bats. All X 2.
Fig. 24. Myotis californicus californicus, no. 18490, 9.
Fig. 25. Myotis californicus pallidus, no. 17787, 2.
Fig. 26. Myotis orinomus, no. 6941, 9.
Fig. 27. Pipistreilus hesperus hesperus, no. 10409, g.
Fig. 28. Myotis longicrus longicrus, no. 20811, 9.
Fig. 29. Myotis yumanensis sociabilis, no. 6651, 2.
Fig. 30. Myotis occultus, no. 10706, ¢.
Fig. 31. Myotis lucifugus altipetens, no, 23036, 9.
Fig. 32. Myotis velifer, no. 7762, 3.
Fig. 33. Lasionycteris noctivagans, no. 24208, 9.
Fig. 34. Myotis evotis, no. 6953, 9.
Fig. 35. Myotis thysanodes, no. 29833 (U. 8. Nat. Mus.), 9.
[ 398 ]
PLATE 22
Side views of the same skulls as shown on plate 21. All X 2.
g-
. 36,
o Bills
. 38.
. 39.
. 40.
ig. 41.
. 42.
. 43.
. 44,
. 45.
. 46.
47.
Myotis californicus californicus, no. 18490, 9.
Myotis californicus pallidus, no. 17787, ¢.
Myotis orinomus, no. 6941, 9.
Pipistrellus hesperus hesperus, no. 10409, JZ.
Myotis longicrus longicrus, no. 20811, 9.
Myotis ywmanensis sociabilis, no. 6651, 9.
Myotis occultus, no. 10706, g.
Myotis lucifugus altipetens, no. 23036, 2.
Myotis velifer, no. 7762, g.
Lasionycteris noctivagans, no. 24208, 2.
Myotis evotis, no. 6953, 9.
Myotis thysanodes, no. 29833 (U. 8. Nat. Mus.), 9.
[ 400 ]
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;
.
NU ¢ 4
PLATE 23
Dorsal views of the skulls of eight species of California bats. All X 2.
Fig. 48. Nyctinomus mexicanus, no. 6948, 3.
Fig. 49. Nycteris cinerea, no. 6944, g.
Fig. 50. Corynorhinus rafinesquii intermedius, no. 6957, 3.
Fig. 51. Nycteris borealis teliotis, no. 24327, 9.—
Fig. 52. EHptesicus fuscus, no. 5176, J.
Fig. 53. Antrozous pacificus, no. 5248, ¢.
Fig. 54. Macrotus californicus, no. 1238, 9.
Fig. 55. <Antrozous pallidus, no. 7346, g.
[ 402 ]
PLATE 24
Side views of the same skulls as shown on plate 23. All X 2.
Fig. 56. Nyctinomus mexicanus, no. 6948, 3.
Fig. 57. Nycteris euierea, no. 6944, ¢.
Fig. 58. Be aha sane intermedius, no. 6957, J.
Fig. 59. Nycteris borealis teliotis, no. 24327, 9.
Fig. 60. Eptesicus fuscus, no. 5176, 3g.
Fig. 61. <Antrozous pacificus, no. 5248, ¢.
Fig. 62. Macrotus californicus, no. 1238, 9.
Fig. 63. <Antrozous pallidus, no. 7346, 2.
[ 404 ]
HO 61
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IN
ZOOLOGY
Vol. 17, No. 13, pp. 405-422, 1 figure in text February 23, 1918
THE PACIFIC COAST JAYS OF THE GENUS
APHELOCOMA
BY
H. 8S. SWARTH
(Contribution from the Museum of Vertebrate Zoology of the University of California)
The present study is based principally upon the series of Aphelo-
coma in the Museum of Vertebrate Zoology, including, in addition
to the specimens contained in the Museum collection, those in the
Grinnell, Morcom and Swarth collections. Besides this material speci-
mens were borrowed from other institutions and from individuals, as
follows: From the Biological Survey, Washington, twelve skins from
the San Pedro Martir Mountains, Lower California: from the Museum
of Comparative Zoology, thirteen from the Cape San Lueas region,
and two from northern Lower California; from the Museum of His-
tory, Science and Art, Los Angeles, four specimens from southern
California ; from the collection of the Oregon Fish and Game Commis-
sion, five specimens from Oregon; from the collection of J. and J. W.
Mailliard, seventeen specimens, mostly from Marin County, Cali-
fornia ; from the collection of J. E. Thayer, twenty-five specimens from
Lower California. The skins borrowed from outside sources are all in
one way or another illustrative of points not covered by the collection
of the Museum of Vertebrate Zoology, and the privilege of utilizing
them added much to the value of the comparisons made. The writer
wishes here to express his appreciation of, and thanks for, these gen-
erous loans. Altogether 437 specimens of Aphelocoma were examined.
Under each form, in the section giving “‘ distinguishing characters,’
I have entered an exact statement of the color of a selected specimen,
an average example of the race in fresh fall plumage. Color terms
used are from Ridgway’s Color Standards and Color Nomenclature
(1912).
406 University of California Publications in Zoology Vou. 17
The genus Aphelocoma is, with the exception of the singularly
isolated Florida species, restricted to the southwestern portion of
North America. The several species included therein are for the
most part rather sharply defined and not prone to split up into local
races, though it is probably true that certain forms now recognized
as “‘species’’ are as yet so imperfectly understood as to leave their
true status and distribution a matter of some doubt.
In California there are three well defined species of the genus,
Aphelocoma californica (divided in this state into three subspecies),
A. woodhousei, and A. insularis. The first mentioned, in its three
forms, occurs over the greater part of the state. It is primarily a
bird of the Upper Sonoran zone, especially favoring those regions
where there is live oak timber or serub oak brush, and extending
locally up into low Transition. It is apparently absent from the
extreme northern coast region, as also from the mountains above
Transition, and from the Lower Sonoran deserts of the southeast.
Aphelocoma woodhousei occupies disconnected areas of Upper
Sonoran on certain of the mountain ranges of the Inyo region, and
on the east slope of the Sierra Nevada. A. insularis is found only on
Santa Cruz Island. These three species are distinguished by certain
trenchant characters. The Santa Cruz Island jay appears to be eut
off from contact with its nearest mainland relative by the intervening
channel, though the distance to be traversed is not so great as, on the
face of it, to be considered an impassable barrier to a bird of this
nature. A. californica and A. woodhousei meet at the east base of
the Sierra Nevada, where at certain seasons both species may be
found at the same places; for the most part the specific characters of
each remain stable along this border line.
The California forms of Aphelocoma all belong to a section of the
genus that it seems to me is deserving of recognition, nomenclaturally,
as distinct from another aggregation now included in the same genus.
Californica, woodhousei, insularis, and hypoleuca, together with cyanea
and certain other Texan and Mexican species, all possess in common
certain conspicuous features of structure, coloration, etc., and all are
as sharply distinguished from the Aphelocoma siebert group. Coues
(1903, pp. 497, 499) proposed that the latter be separated as a sub-
genus ‘‘Sieberocitta,’’ as distinguished from true Aphelocoma. This
suggested subgenus was not aecepted by the A. O. U. Committee (1908,
p. 394), because of being ‘‘based on color characters alone.’’ As
a matter of fact the distinguishing features (as was pointed out by
1918] Swarth: The Pacific Coast Jays of the Genus Aphelocoma 407
Coues) are matters of structure (at least such as are commonly used
in distinguishing bird genera), of color, of eggs, and of habits. In
Sieberocitta, as compared with true Aphelocoma, besides the obvious
and constant color differences, the tail is shorter instead of longer
than the wing, and the eggs are plain blue instead of double pigmented.
The call-notes of the two are widely different. Of habit differences
it may be noted that while Aphelocoma is relatively solitary in its
mode of life, Sieberocitta is as markedly gregarious as the Pinon
Jay (Cyanocephalus cyanocephalus), going habitually in closely
assembled flocks of twenty or thirty individuals, and even nesting in
rather loose rookeries. Another feature of Sieberocitta is its habit
of constructing extra nests, as the Marsh Wren does.
The point here made is that in the genus Aphelocoma there are
these two aggregations of species and subspecies, each group dis-
tinguished by certain features, common to all the forms therein, of
coloration and pattern, structure, eggs, call-notes and habits. The
two groups occur to a large extent over the same territory, where,
however, they occupy slightly different ecologic niches, Sieberocitta
being primarily a bird of open oak woods, true Aphelocoma, of denser
underbrush. It seems to me that it is desirable that these two groups
be accorded subgenerie recognition.
If it were feasible to consider all the forms of true Aphelocoma as
subspecies of one species, and all of Sicberocitta as of another, this
might equally well serve to segregate them as they should be, but
the facts hardly permit of such treatment.
There is still another feature of the situation, namely, the relation-
ship of the Mexican and Central American species, Aphelocoma wni-
color; but this I am not qualified to discuss.
In a recent paper, primarily on the status of Aphelocoma texana,
but incidentally discussing other allied forms, Oberholser (1917, p.
94) advocates that all of the ‘‘Aphelocoma californica group’’ be
regarded as subspecies of one species, A. californica. This includes
all the forms that I would restrict to the subgenus Aphelocoma,
except A. cyanea and A. insularis. Mr. Oberholser and myself are,
perhaps, merely stating the same thing in different ways. My main
objection to his conception of the forms as subspecific variants is
that much of the intergradation to which he calls attention occurs
only through individual variation, in races that are geographically
far apart. Individual variation in swmichrasti, in southern Mexico,
is said to cover the difference between that form and A. californica
408 University of Califorma Publications in Zoology [ Vou. 17
californica (Oberholser, loc. cit.), and the same is true as between
A. californica immanis of the Sacramento Valley, and A. hypoleuca
of Cape San Lueas, but in each case the region between these forms
is occupied by a race or races different from either.
It seems to me that there is danger of giving too much emphasis
to the resemblances noted, and of losing sight of the significant fact
that there is no blending of characters where the different forms
meet. For example, although in the three subspecies of Aphelocoma
californica, as I would restrict them, californica, oocleptica, and
immanis, there is such intergradation at the margins of the several
habitats, nothing of the sort can be detected along the boundary
between immanis and woodhouset. It is true that there is a specimen
at hand that may be regarded as intermediate between these two
latter forms (see beyond), but this is a single sporadic individual, and
the circumstance appears to be exactly comparable with conditions
observed to exist in Psaltriparus minimus californicus and P. plum-
beus, of the same region. In both cases in this marginal region the
populations in general of the several species are as typical in appear-
ance as are those at opposite extremes in the birds’ ranges. It is
just an occasional individual which shows any admixture of characters
of the adjoining races.
The manner of variation to be traced through the several sub-
species of Aphelocoma californica differs in some respects from what
may be observed among other variable species of birds. Aphelocoma
is not an especially ‘‘plastic’’ group, the several species being for
the most part rather sharply defined, and remaining uniform in
appearance over a wide expanse of territory. In the three recogniz-
able subspecies of A. californica occurring within the state of Cali-
fornia, the manner of variation is unlike what is observed in the races
of such species as Pipilo maculatus and Thryomanes bewicki, for, as
shown in the accompanying table, in A phelocoma the several characters
concerned vary, to all appearances, independently, and not always in
the same direction.
Characters which have been used to differentiate species and sub-
species in the genus Aphelocoma pertain to size and color. Size char-
acters consist of general size, length of wing, length of tail, length of
bill and bulk of bill. Color characters used are: shade of the blue
areas, of dorsum, of under parts, of under tail coverts.
It should be noted that characters of color are quite as apparent
in the juvenal plumage as in the adult, and, when the quill feathers
Color of under surface of body
Color of under tail coverts
Juvenal plumage
Cape San Lucas
(A. hypoleuca)
Pale Neropalin
blue
Light drab, suffused
with bluish
White, light suf-
fusion of gray
Variable features in Aphelocoma californica and A. hypoleuca listed to show differences occurring in the several faunal areas occupied.
Northern Lower
California
(A, ce. californica)
San Diegan region
(A. ¢. californica)
Santa Cruz region
(A. c. californica)
San Francisco
Bay region
(A. c. oocleptica)
Coast region north of
the Golden Gate
(A. ¢. oocleptica)
Sacramento Valley
(A. ¢. immanis)
Sierra Nevada
(A. ¢. immanis)
Warner Mountains
(A. ¢. immanis)
Oregon
(A. c. immanis)
Smaller Smaller Smaller Larger Larger Larger
120.4 118.8 119.8 126. 125.4 126.4
134.4 132.5 131.6 140. 136.7 137.7
8.3 9.1 9.5 9.1 9.7
Dark (deep cadet
blue)
Dark (deep cadet
blue)
Dark (deep cadet
blue)
Dark (deep cadet
blue)
Dark (deep cadet
blue)
Sepia
Slaty
White, frequently
tinged with blue
Sepia
Sepia
Sepia
Sepia
Chapman’s blue
Drab
Slaty
Slaty
Slaty
Whitish with slaty
suffusion
Chapman’s blue
Drab
Chapman’s blue
Drab
White, light suf-
fusion of gray
White, frequently
tinged with blue
White, frequently
tinged with blue
White, frequently
tinged with blue
White
Dark
Dark
Dark
Dark
White, light suf-
fusion of gray
White, light suf-
fusion of gray
Chapman’s blue
Drab
White, light suf-
fusion of gray
White
Pale
408 University of California Publ
californica. (Oherholser_ loc. cit.). and
A. califoi
of Cape }
is oceuple
It see
to the res
that ther
meet. F\
californic
immanis, .°
habitats,
between 2
at hand
latter for e} Sele ef:
the circu
observed
beus, of
populatic
ance as
just an o:
of the ad
The 1
species 0
may be ¢
is not al
the most
appearal
able sub;
fornia, t]
of such ;
shown in
concerne
the same
Char:
species
acters ¢c
bill and
areas, of
Tt shove Se Se
in the juvenal plumage as in the adult, and; when
1918] Swarth: The Pacific Coast Jays of the Genus Aphelocoma 409
have attained their full size, length of wing and tail are also equally
characteristic in the young. In several instances in the present study,
where localities have been represented by adults in worn, midsummer
plumage, the appearance of young birds taken at the same time
pointed unmistakably toward the relationships of the series in
question.
On the accompanying table the variable features in Aphelocoma
californica are listed so as to show the differences occurring in the
birds of the several faunal areas inhabited. Where there is little or
no difference in a character in two or more of such areas, the dividing
line is lightly indicated. Where there is an appreciable discrepancy
this line is heavy. Thus in the diagram, differentiating characters
are segregated by locality regardless of subspecifie names.
It will be observed of the Cape San Lucas bird (hypoleuca) that
in every particular it is sharply set off from its nearest neighbor,
californica, so much so that, if, as seems highly probable, there is a
stretch of country between the two uninhabited by the genus, cali-
fornica and hypoleuca might well be considered distinct species rather
than subspecies of the same form. It is true that although Aphelo-
coma hypoleuca and A. californica as occurring in northern Lower
California (A. c. californica) are so abruptly and absolutely different,
hypoleuca and Sacramento valley californica (A. c. immanis) are dis-
tinguished with difficulty ; but this seems to be a case of parallel modi-
fication of widely separated forms, races that really have but little in
common.
It will also be noted upon the diagram, relating to the species as
occurring in California and Oregon, that although no line of separa-
tion can be drawn between any two adjacent regions for all the char-
acters considered, certain divisions can be made by combination of
various of the differentiating features. Disregarding the Cape San
Lucas bird, it will be seen that beginning at northern Lower Cali-
fornia there is increase of size northward; not evenly, however, for
although a line drawn between the Santa Cruz and San Francisco
Bay regions will separate smaller sized birds to the southward from
larger ones to the northward of this point, there is no appreciable
size difference between birds of the Santa Cruz area and northern
Lower California on the one hand, nor between those of the San
Francisco Bay area and Oregon, on the other.
There is no appreciable difference in length of culmen from Cape
San Lucas to Oregon (see table), but depth of bill increases north-
410 University of California Publications in Zoology [Vou. 17
ward, more northern birds consequently possessing proportionately
heavier bills. As seen in the diagram it is in this one particular that
birds of the Santa Cruz area resemble the northern form more nearly
than the southern.
In coloration it will be noted that there is practical uniformity
of appearance, on the one hand, in birds of the coast region from
northern Lower California to Humboldt Bay, on the other, in birds
from the Sacramento-San Joaquin Valley, Sierra Nevada, Warner
Mountains, and Oregon.
Aphelocoma californica californica (Vigors)
Type locality —Monterey, California.
Range.—A relatively narrow strip along the coast of California and northern
Lower California; from the San Pedro Martir Mountains, Lower California,
north on the coastal slope of California, west of the southern Sierras and the
coast ranges, through the San Diegan and Santa Cruz districts to the south
side of San Francisco Bay.
Specimens examined from the following localities: Lower California: San
Pedro Martir Mountains, 6; Hanson Laguna Mountains, 5; Ensenada, 1; 30
miles east of San Quintin, 1; Santana, 2.
California: San Diego County—Witch Creek, 7; Cuyamaca Mountains, 2;
Julian, 4; Foster, 2; Dulzura, 1; Campo, 1; Warner Pass, 1; San Diego, 1;
Pala, 1. Orange County—Trabuco Cafion, 2; Laguna Beach, 1. Riverside
County—San Jacinto Mountains, 16; Vallevista, 1; San Gorgonio Pass, 2;
Riverside, 1. San Bernardino County—Reche Canon, 3; San Bernardino Moun-
tains, 5. Los Angeles County—Pasadena, 39; San Fernando Valley, 2; Santa
Monica Mountains, 4; Verdugo, 1; Glendora, 2; Chileo, 1; San Francisquito
Cafion, 1; Los Angeles, 3. Ventura County—Ventura, 2; Mount Pinos, 3.
Santa Barbara County—Santa Barbara, 1; Guadalupe Lake, 1. San Luis
Obispo County—Paso Robles, 1. San Benito County—Paicines, 2. Monterey
County—Pacifie Grove, 3; Partington Pt., 2; Lucia 2. Santa Clara County—
Palo Alto, 7; San Jose, 1; Berryessa, 1; College Park, 2. Total 147.
Distinguishing characters.—Aphelocoma californica californica, compared with
A. c. immanis, is of small size and dark coloration. The blue areas are of a
deeper shade, the back distinctly darker brown, and the light colored under
parts have a dusky suffusion. Lower tail coverts usually tinged with blue,
sometimes conspicuously so. Coloration is about the same in californica as in
oocleptica, from which subspecies californica is distinguished by smaller size
throughout. Compared with A. hypoleuca, A. ec. californica is of much darker
coloration, and, on the average, slightly smaller size. No. 7016 (Mus. Vert.
Zool.), male, Palo Alto, California, November 21, 1906. Blue areas, deep cadet
blue; dorsum, sepia.
Remarks.—In the material assembled for the present study the
subspecies Aphelocoma c. californica is represented by a fairly satis-
factory series from Lower California (unsatisfactory only in that it
includes no specimens in fresh fall plumage), by skins from the San
1918]
Aphelocoma
Aphelocoma
Aphelocoma
Aphelocoma
Aphelocoma
Aphelocoma
eor1e¢e+ > 8
Fig. 1. Distribution of the species and sub-
species of Aphelocoma on the Pacifie Coast.
Swarth: The Pacific Coast Jays of the Genus Aphelocoma
Points from which specimens were examined:
californica californica
californica immanis
californica oocleptica
woodhousei
insularis
hypoleuca
Cape San Lucas
411
412 University of California Publications in Zoology [Vou. 17
Diegan region representing a large number of record stations and
illustrating practically all phases of variation, and by smaller series
from a number of points along the coast north to San Francisco Bay.
The most important point to be considered in a systematic study
of birds from these several sections is the relationship borne by
A. c. obscura, described from the San Pedro Martir Mountains, Lower
California, to A. c. californica, with type locality at Monterey. The
present treatment of the races of the California jay differs from that
in most recent literature covering the subject (e.g., A. O. U. Check-list,
1910, p. 225; Ridgway, 1904, pp. 827-331) in that it does not recognize
the subspecies obscura. This race was described by Anthony (1889,
p. 75) from specimens taken in the San Pedro Martir Mountains,
Lower California. In a subsequent paper (1893, p. 239) the same
writer asserts that birds from the San Pedro Martir Mountains and
from San Diego County, California, are indistinguishable, and for
some years past the name obscura has been generally used to cover
the bird of the San Diegan region of California, as well as that of
northern Lower California. Comparison of series from these points,
however, with specimens from various coastal localities as far north
as San Francisco Bay (including the vicinity of Monterey, the type
locality of californica), shows that all belong to the same race, that
there are no characters serving to distinguish specimens from these
several places. Hence the name obscura must be considered a synonym
of californica.
Aphelocoma californica obscura was described as a smaller and
darker colored bird than A. c. californica. Perpetuation of this error
may have occurred through comparison of southern California speci-
mens with others from the Sacramento Valley or the Sierra Nevada,
in the belief that the latter were representative of typical californica.
This assumption is wrong, however, and although jays from certain
sections of California may readily be distinguished as, respectively,
larger and paler, or smaller and darker, true californica and obscura
both fall into the latter category. This point has been discussed and
the same conclusion reached by Grinnell and Swarth (1913, p. 261),
with reference to the status of the species as occurring in the San
Jacinto Mountains, southern California.
The range of Aphelocoma c. californica abuts that of A. c. ooclep-
tica on the north, and that of A. c. immanis on the northeast. Whether
or not there is continuous distribution of jays of this species over the
length of Lower California, with consequent contact of the ranges of
1918] Swarth: The Pacific Coast Jays of the Genus Aphelocoma 413
californica and hypoleuca, I do not know. There are no specimens of
either at hand illustrative of marked variation toward the neighboring
race.
In southern California, as far north as Fort Tejon, the eastern
boundary of the range of californica is sharply defined by the western
margins of the Colorado and Mohave deserts. At this point the sub-
species califormca extends in typical form to the eastern bases of the
surrounding coast ranges, and stops there abruptly. The Lower
Sonoran deserts form a broad and impassable barrier to birds of this
genus.
Farther north, from the northern boundary of Ventura County
northward, while there is no such evident obstacle to distribution,
there is an extensive stretch of country, the arid, timberless west side
of the San Joaquin Valley, where conditions are so unfavorable to
the species that it is almost, if not entirely, absent. I have considered
the range of A. c. californica as lying just west of this section. There
are no specimens at hand from any point in this region. There are
available three skins from the vicinity of Mount Pinos, Ventura
County, and two from Paicines, San Benito County, these points
lying approximately along the dividing line between the ranges of
californica and immanis. Of the specimens from the Mount Pinos
region, two November birds from the head of Piru Creek show a
decided approach to Sierra Nevada immanis. In fact, regarded by
themselves they might well be considered as belonging to that sub-
species, but considering the manner in which the seetion where they
were taken is separated from the range of immanis, and the econtin-
uous distribution of californica from this point westward, they had,
perhaps, better be regarded as individual extremes of californica,
taken at the edge of its range and illustrating intergradation with
immanis. The two Paicines specimens are also evidently intergrades
toward immanis, though not leaning so markedly toward the latter
race. There are specimens at hand from various points along the
coastal slope of Santa Barbara, San Luis Obispo, Monterey, and
Santa Clara counties, as already enumerated, and these are so
obviously like the bird of southern California and northern Lower
California as to leave no doubt as to their subspecifie identity.
Aphelocoma californica oocleptica, new subspecies
Type.——Male adult, no. 7123, Mus. Vert. Zool.; Nicasio, Marin County,
California; February 23, 1909; collected by Walter P. Taylor; original num-
ber 647.
414 University of California Publications in Zoology [Vou. 17
Range.—The coast region of northern California, west from Mount Diablo
and the coast ranges. North to Humboldt Bay, south to the Golden Gate and
the east side of San Francisco Bay.
Specimens examined from the following localities: Alameda County—Oak-
land, 1; Piedmont, 1; Berkeley, 9. Contra Costa County—Walnut Creek, 5;
Mount Diablo, 3; Danville, 5. Marin County—Mailliard, 1; Inverness, 3;
Nicasio, 4; San Geronimo, 3. Sonoma County—Sonoma, 1; Stony Point, 2; Free-
stone, 1; Guerneville, 10; Santa Rosa, 1. Mendocino County—Mendocino City, 1;
Mount Sanhedrin, 2; Bald Hill, 1; Sherwoods, 2. Humboldt County—Arcata, 1.
Total 57.
Distinguishing characters.—Of large size and dark coloration. In color closely
similar to A. ec. californica, but size measurably greater throughout. In measure-
ments oocleptica is equal to the maximum of immanis, from which subspecies it
is distinguished by its dark coloration. Differs from hypoleuca both in greater
size and much darker color.
No. 6001 (Mailliard coll.), male, Nicasio, Marin County, October 29, 1894.
Blue areas, deep cadet blue; dorsum, sepia.
Remarks—It was an unexpected development of the present study
that there should be disclosed the presence of an additional race of
Aphelocoma within the state. The specimens at hand, however, are
ample for the establishing of this subspecies, the characters exhibited
being clearly defined, and the territory occupied by the form capable
of being outlined with a fair degree of accuracy. While in the com-
bination of characters possessed oocleptica might be considered as
an intergrade between californica and immanis, still it is not truly
intermediate between the two. It has assumed the distinetive char-
acters of large size (equal to the pale colored immanis), and dark
eoloration (as in the small sized californica), each to the fullest degree.
It is the combination of these two features that distinguishes the race.
There is more evident appearance of intergradation between
oocleptica and immanis, than between the former and californica.
Of the last mentioned form, examples from points immediately south
of San Francisco Bay (closely approaching the range of oocleptica)
are indistinguishable from specimens from southern California, there
being no appreciable increase in size. On the other hand, at points
where the ranges of oocleptica and immanis come together (as in
parts of Marin and Sonoma counties), individuals exhibiting various
degrees of intergradation between the two are of frequent occurrence.
In southern Marin County somewhat similar conditions prevail in
Aphelocoma as have already been noted of other variable groups in
the same region as Psaltriparus (Swarth, 1914, pp. 5138-515) and
Thryomanes (Swarth, 1916, p. 66), namely, the frequent occurrence
of individuals evincing a marked tendency toward the appearance of
1918] Swarth: The Pacific Coast Jays of the Genus Aphelocoma 415
the neighboring race of the nearby Sacramento Valley. There is a
series of Aphelocoma at hand from that portion of Sonoma County
wherein, as deseribed by J. Mailliard (1908, p. 133), there oceurs a
form of Cyanocitta closely approaching the interior form frontalis
rather than the coastal race carbonacea. In the Aphelocoma of this
strip of country it is again of interest to note that in birds taken at
points extending practically to the coast, coloration is appreciably
paler than in typical oocleptica, closely approaching that seen in
Sacramento Valley immanis.
In the region immediately west of San Francisco Bay, birds from
the vicinity of Berkeley and Oakland exhibit the extreme of large
size and dark coloration. Just a few miles away, however, on the
farther slope of the hills to the westward of these points, the jays
exhibit a pronounced leaning toward immanis. <A series of five birds
taken at a point in this region (near Danville, Contra Costa County),
and in fresh fall plumage, are noticeably pale colored, strikingly
similar to specimens from the Sierra Nevada. I include this region
in the range of oocleptica, however, partly because of the slightly
darker tone of coloration of adult birds from this section (as com-
pared with Sierran immanis), and largely because in the juvenal
plumage (as exhibited in specimens from Mount Diablo and Walnut
Creek) there is no variation from the corresponding stage as it occurs
in Marin County. In typical immanis the juvenal plumage is as
conspicuously different from oocleptica as is the adult.
Aphelocoma californica immanis Grinnell
Type locality.—Scio, Linn County, Oregon.
Range.—Extreme southern Washington, in Oregon those valleys lying between
the Cascades and the Coast Ranges, and south in California through the Sac-
ramento and San Joaquin valleys and the Sierra Nevada. East to the Warner
Mountains and the eastern base of the Sierra Nevada.
Specimens examined from the following localities:
Oregon: Linn County—Scio, 4. Benton County—Corvallis, 1. Multnomah
County—Portland, 2. Josephine County—Grant’s Pass, 1. Jackson County—
Gold Hill, 2. Douglas County—Roseburg, 1. ‘‘ Willamette Valley,’’ 1.
California: Modoe County—Warner Mountains, 38. Trinity County—Helena,
1. Shasta County—Baird, 1. Nevada County—Independence Lake, 1; Little
Truckee River, 1. Placer County—Cisco, 1. Inyo County—Kearsarge Pass, 2;
Carroll Creek, 1. Kern County—Onyx, 1; Walker Pass, 1; Fay Creek, 5; Bod-
fish, 2; Piute Mountains, 4; Caliente, 1. Tulare County—Taylor Meadow, 4;
Trout Creek, 4; Cannell Meadow, 2. Fresno County—Dunlap, 4; Minkler, 14.
Madera County—Raymond, 1. Mariposa County—El Portal, 7; Coulterville, 2.
Merced County—Snelling, 2. Stanislaus County—La Grange, 1; Modesto, 3.
416 University of California Publications in Zoology [ Vou. 17
San Joaquin County—Tracy Lake, 3. Sacramento County—Sacramento, 3.
Amador County—Carbondale, 2; Drytown, 1. Solano County—Vaeaville, 5.
Glenn County—Winslow, 5. Yolo County—Knight’s Landing, 1. Butte County
—Oroville, 1. Sutter County—Sutter, 1. Lake County—Lower Lake, 1. Total
139.
Distinguishing characters.—Of large size and pale coloration. Under tail
coverts usually pure white; sometimes slightly tinged with blue; brown of
back pale, as compared with californica, and suffused with grayish. Under parts
white, with but a slight suffusion of slaty. Distinguished from 4. c. californica
both by large size and pale coloration; from oocleptica by pale coloration, size
being about the same. Aphelocoma hypoleuca is smaller than the maximum of
immanis, though closely matched in this respect by the series from the Sacra-
mento Valley. In coloration hypoleuca is constantly paler than the lightest
colored immanis.
No. 269 (coll. of S. G. Jewett), male, Portland, Oregon, November 5, 1905;
blue areas, Chapman’s blue; dorsum, hair brown, with bluish tips to the feathers.
Remarks—Aphelocoma californica tmmanis was described by
Grinnell (1901, p. 188), from the Willamette Valley, Oregon, char-
acterized as a bird of larger size and with longer tail than A. c. cali-
fornica. The subspecies was denied recognition by the A. O. U. Com-
mittee (1901, p. 312), as indistinguishable from A. c. californica, a
logical point of view, of course, considering the recognition already
accorded by that committee to A. c. obscura as a smaller and darker
bird than true californica. There is no question as to the existence of
the two distinguishable races, the large, pale colored form to which
the term californica is incorrectly applied by Ridgway (1904, p. 327)
and the A. O. U. Committee (1910, p. 225), and the small, dark colored
subspecies termed obscura by the same authorities. The realization,
however, that obscura is a synonym of californica, both pertaining
to the subspecies occurring in the southern coast district of California,
necessitates the affixing of a name to the form inhabiting the interior
of the state. In a recent paper Oberholser (1917, p. 94) affirms the
existence of the subspecies immanis, defining its range as including
parts of Oregon and extreme northern California. There are at hand
large series of jays from the interior valleys of California, the Sierra
Nevada, and the Warner Mountains, and as, on comparison, these are
indistinguishable from specimens from the Willamette Valley, Oregon,
the name which has been used for the latter bird, immanis, must be
applied to this whole aggregation.
On the east slope of the Sierra Nevada the ranges of immanis and
woodhousei meet, and here is where intergradation of characters
between the two should be found if it occurs at all. Woodhousei is
found in this section in the fall, but whether or not it breeds here is
1918] Swarth: The Pacific Coast Jays of the Genus Aphelocoma 417
not known. The probabilities are that it does not, and that the birds
that have been taken in the region in the autumn were wanderers from
points farther east. However this may be, there is one specimen at
hand from this section, taken at 5,500 feet elevation on Carroll Creek,
Inyo County, that in color and markings exhibits a blending of the
characters of the two forms. This bird, an adult male (no. 20068,
September 9, 1911), is of a decidedly paler blue than are comparable
specimens from the west side of the Sierras, while the dorsum is
bluish gray, rather than brownish, as in the latter. Whiteness of
under parts of body and lower tail coverts, the streakings upon the
breast, and measurements and proportions, are all exactly as in
immanis. This bird therefore must be classed with the latter form,
but the dorsal appearance of the specimen is strikingly similar to fall
examples of woodhousei, and quite different from any other specimen
of immanis at hand. The nature of this departure from the usual
coloration of immanis, occurring in a specimen from this particular
region has undoubtedly some significance as regards the relationships
of the two forms. For the present, however, I prefer to regard them
as specifically distinet, for the same reasons as those advanced in the
ease of the two species of Psaltriparus that occur here; the observed
conditions are closely similar. (See Swarth, 1914, p. 521.)
There are two breeding birds at hand from this section, taken near
the east base of Kearsarge Pass in June (nos. 22504, 22505). The
feathers are too abraded to be of service in color comparisons, but to
all appearances these birds are typical of ammanis.
Aphelocoma woodhousei (Baird)
Type locality.—Fort Thorn, New Mexico.
Range in California—Upper Sonoran zone in the desert mountains of the
eastern part of the state, in the Inyo and Mohave regions. At the eastern
base of the Sierra Nevada, probably as a transient only.
Specimens examined from the following localities:
California: Inyo County— Carroll Creek, 1; Hanaupah Cafion, Panamint
Mountains, 3; Johnson Caton, Panamint Mountains, 6; Jackass Spring, Pana-
mint Mountains, 9; Silver Cafion, White Mountains, 6; Robert’s Ranch, White
Mountains, 1; Keeler, 1. Mono County—Williams Butte, 2; Benton, 3.
Arizona: Huachuea Mountains, 5. Santa Catalina Mountains, 2. Rincon
Mountains, 1. Dragoon Mountains, 3.
Nevada: Quinn River Crossing, Humboldt County, 1.
Total 44. .
Distinguishing characters—Compared with any of the subspecies of Aphelo-
coma californica, A. woodhousei differs in coloration and in proportions of bill.
The blue areas are dull and pale, the back is strongly suffused with bluish gray,
418 University of California Publications in Zoology [Vou. 17
and the under parts and throat with gray; the under tail coverts are blue.
The general effect of these modifications is to produce a much more uniformly
and inconspicuously marked bird than 4A. californica. The bill of woodhousei
averages longer than in californica, but is more slender. From 4A. insularis,
woodhousei differs constantly in its subdued and uniform coloration, and smaller
size.
No. 25934 (Mus. Vert. Zool.), male, Williams Butte, Mono County, Cali-
fornia, September 21, 1915. Blue areas, Columbia blue; dorsum and lower
parts, suffused with bluish throughout.
Remarks.—The range of the Woodhouse jay in California is re-
stricted to scattered and disconnected areas of Upper Sonoran in the
Inyo region, the arid desert section of the eastern part of the state.
In the late summer and fall it is a visitant to the eastern slope of the
Sierra Nevada, where it comes into direct contact with A. c. immanis,
but it apparently does not breed in this section. Carroll Creek, just
south of Mount Whitney, is the southernmost record station, though
there is no evident reason why the species should not extend much
farther south and west.
Comparison of three California specimens at hand in fresh fall
plumage, with individuals taken at the same season in southern
Arizona, shows no differences between the two lots.
Aphelocoma insularis Henshaw
Type locality—Santa Cruz Island, California.
Range.—Santa Cruz Island, California.
Specimens examined.— From Santa Cruz Island, 14.
Distinguishing characters.—Distinguished from any subspecies of A. cali-
fornica by greater size, darker coloration, and definitely blue under tail coverts.
From A. woodhousei, which it resembles in its blue under tail coverts, insularis
is distinguished by greater size, darker coloration, and (like A. californica)
in more strongly contrasted markings.
No. 5458 (coll. of J. Grinnell), male, Santa Cruz Island, September 3, 1903.
Blue areas, deep dull violaceous blue; dorsum, clove brown.
Remarks.—The Santa Cruz jay is in many respects one of the most
remarkable species that has been produced upon the Santa Barbara
Islands. Each of the animal forms evolved upon these islands has
developed in ways deserving the most careful study, for neither in
the trend taken by the characters serving to distinguish them from
their mainland relatives, nor in the distribution of species upon the
islands, is it possible to detect uniformity in the results attained. In
the particular species in question certain anomalies stand out even
more conspicuously than in any of the other island birds.
1918] Swarth: The Pacific Coast Jays of the Genus Aphelocoma 419
The Santa Cruz jay is one of the most sharply differentiated of
any of the island species, and it is hard to appreciate the possibility
of the development of the form under the given conditions. Santa
Cruz is the only one of the Santa Barbara Islands upon which jays
of any sort are found, though other species of birds far more sedentary
in their manner of life, are widely distributed over the group, and
conditions upon some, at least, of the other islands would permit the
existence of the jay. Santa Cruz is not so widely distant from the
mainland but what it would seem possible for jays to travel back and
forth. While the California jay of the adjacent mainland strictly
speaking is not a migratory species, still in the fall small flocks may
oceasionally be seen wandering far from their breeding grounds, in
territory where they certainly never nest, and it would seem not
unlikely, under the circumstances, for individuals from the mainland
to reach the islands occasionally, and vice versa.
Then, as regards the extreme differentiation achieved by the
Santa Cruz jay, it is of interest to make comparisons with other
variable forms occurring upon the same island.
The spotted towhee (Pipilo maculatus) as it occurs upon Santa
Cruz is indistinguishable from the mainland subspecies, megalonyx,
though upon others of the islands there is the subspecies clementae,
with strongly developed characters. The Bewick wren (Thryomanes
bewicki) upon Santa Cruz Island is recognized separately, as the
subspecies nesophilus, but it is with difficulty distinguished from the
race charienturus upon the adjoming mainland, while again upon
other of the islands there are races with much more strongly marked
features. The bush-tit (Psaltriparus minimus), Hutton vireo (Vireo
huttont), and perhaps one or two other species, occur upon Santa
Cruz alone of the Santa Barbara Islands—as does the jay—yet none
of these have developed into recognized insular races.
On the whole it is evident that Santa Cruz Island has not served
as a differentiating center to the same extent as other islands of the
group; yet at the same time it has produced in the Santa Cruz jay
the most strongly characterized of any of the island races.
The most striking feature of the Santa Cruz jay, as compared with
the mainland species is its enormous size, so in this case a marked
restriction of range, with consequent probability of inbreeding of
closely related individuals has not been productive of the dwarfed
stature which such conditions are supposed to engender.
420 University of California Publications in Zoology [ Vou. 17
Aphelocoma hypoleuca Ridgway
Type locality.—La Paz, Lower California.
Range—‘ ‘Cape district of Lower California ..., north to about latitude
28°’? (Ridgway, Birds of North and Middle America, 3, 1904, p. 331).
Specimens examined from the following localities:
Lower California—La Paz, 8; Miraflores, 10; El Sanz, 4; Sierra de Laguna,
5; Santa Anita, 1; Cape San Lucas, 7; Triunfo, 1.
Total 36.
Distinguishing characters —The pale coloration of hypoleuca serves to dis-
tinguish it from any of the related species or subspecies on the Pacifie Coast.
The blue areas are distinctly lighter than in even the palest examples of
immanis, and the under parts are of a more nearly immaculate white.
No. 11917 (coll. of John E. Thayer), male, Sierra de Laguna, Lower Cali
fornia, August 25, 1908. Blue areas, Neropalin blue; dorsum, light drab, suf-
fused with bluish.
Remarks.—It is a difficult matter to come to a decision as to the
proper nomenclatural treatment that should be accorded this form,
though it is my impression that it is best regarded as a distinet species,
rather than as a subspecies of Aphelocoma californica. The one great
objection to this course is the close resemblance of hypoleuca to the
form of Aphelocoma occurring in northern California. Whether this
possibly fortuitous general resemblance of the two is sufficient to out-
weigh the several arguments in favor of their distinctness, is a ques-
tion, but the fact remains that it is almost impossible to designate
characters whereby hypolewca and series of immanis from certain
points can be distinguished without fail.
So much may be said in opposition to the concept of specific differ-
ence. The arguments in favor of such a view are essentially as follows:
1. The features in which hypoleuca differs from the form geo-
graphically nearest to it (A. c. californica) are not in accord with
the trend of variation observed in A. californica over its range as
a whole.
2. Between californica of northern Lower California, and hypo-
leuca of the southern end of the peninsula, the change in appearance
is abrupt, comparable to the difference between immanis.and wood-
housei, where the latter meet in California. Two specimens of cali-
fornica from Santana, Lower California (Thayer coll. nos. 6353, 6552)
are indistinguishable from comparable examples from southern Cali-
fornia. Santana is about midway the length of Lower California,
and at a point where, if intergradation between californica and hypo-
leuca occurs, specimens secured should illustrate this condition. That
Bee
1918] Swarth: The Pacific Coast Jays of the Genus Aphelocoma 421
these two skins are typical of californica is an argument in favor of
the specific distinction of the two forms.
As regards the first of these two points, it will be noted that
although series of californica from northern California (A. c. im-
manis), are closely similar to hypoleuca, the form geographically
closest adjacent to the range of hypoleuca ( A. c. californica, of the
San Pedro Martir Mountains) is the farthest removed from it in all
its characters.
As can be seen on the accompanying table (opp. p. 408) the course
of variation in Aphelocoma californica is, in general terms, as follows.
At the northern extreme of its range there is a large, pale colored
form, restricted to the interior valleys and mountains. This, in the
central coast region merges into a large, dark colored race, which,
in turn, to the southward passes into the smaller, but dark colored,
race that extends into northern Lower California. Then, abruptly,
after what appears to be an absolute break in continuity of range,
there is the large, pale colored form, hypoleuca.
The fact that hypoleuca has been heretofore described as smaller
than californica is probably due to comparisons being made with
series of the form ammanis. Compared with A. c. californica from the
Santa Cruz or San Diegan districts it is, as shown in the accompany-
ing table, of larger size than these birds. Measurements of hypoleuca
are about equal to those of immanis from the Sacramento Vallev or
Sierra Nevada, and smaller than those of series from extreme northern
California and Oregon.
Transmitted June 12, 1917.
422 University of California Publications in Zoology [Vou 17
LITERATURE CITED
AMERICAN ORNITHOLOGISTS’ UNION CoMMITTEE; J. A. ALLEN, CHAIRMAN AND
EDITOR
1901. Tenth supplement to the American Ornithologists’ Union Check-list of
North American birds. Auk, 18, 295-320.
1908. Fourteenth supplement. ... Auk, 25, 343-399.
1910. Check-list of North American birds. Ed. 3, revised (New York;
American Ornithologists’ Union), 430 pp., 2 maps.
ANTHONY, A. W.
1889. New birds from Lower California, Mexico. Proc. Calif. Acad. Sci.,
(2), 11, 73-82.
1893. Birds of San Pedro Martir, Lower California. Zoe, 4, 228-247.
Cougs, E.
1903. Key to North American birds. Ed. 5, revised (Boston: Dana, Estes
and Company), 1, pp. xli, 1-535, 2 pls., figs. 1-353 in text; 2, pp. vi,
537-1152, 1 pl., figs. 354-747 in text.
GRINNELL, J.
1901. The long-tailed jay. Auk, 18, 188.
GRINNELL, J., AND SWARTH, H. S.
1913. An account of the birds and mammals of the San Jacinto area of
southern California, with remarks upon the behavior of geographic
races on the margins of their habitats. Univ. Calif. Publ. Zool., 10,
197-406, pls. 6-10.
MAILLIARD, J.
1908. Sierra forms on the coast of Sonoma County, California. Condor, 10,
133-135, 2 figs. in text.
OBERHOLSER, H. U.
1917. The status of Aphelocoma cyanotis and its allies. Condor, 19, 94-95.
Ripeway, R.
1904. The birds of North and Middle America. U.S. Nat. Mus. Bull., 50,
part 3, xx + 801 pp., 19 pls.
Swartu, H. S.
1914. The California forms of the genus Psaltriparus. Auk, 31, 499-526,
1 pl. (map).
1916. The Pacific coast races of the Bewick Wren. Proce. Calif. Acad. Sci.
(4), 6, 53-85, 1 pl. (map).
Wing
Aphelocoma hypoleuca
10 males, Cape region, Lower CENT 1 2 i a a a 124.2 (120.5-127.0)
Aphelocoma californica californica
7 males, San Pedro Martir and Hanson Laguna Mts.,
Te RN 120.4 (117.0-125.0)
6 males, San Diego and Orange counties, California __. ---- 118.8 (114.0-122.0)
10 males, Los Angeles County, California 119.0 (116.0-121.0)
8 males, Monterey and Santa Clara Counties, California _.... 119.8 (118.0-122.0)
Aphelocoma californica oocleptica
T males, Marin County, California 2200.00. 125.4 (122.0-128.0)
Aphelocoma californica immanis
8 males, Sacramento Valley, California ......... 124.3 (122.0-129.0)
10 males, Sierra Nevada, California 126.0 (122.0-128.0)
6 males, Warner Mts., California 127.0 (125.0-134.0)
EE aS ee a ea 126.4 (121.0-131.0)
Aphelocoma insularis
6 males, Santa Cruz Island, California 135.5 (134.0-138.0)
Aphelocoma woodhousei
‘males, California and Arizona ........ 125.8 (121.0-129.0)
Tail
138.2 (133.0-146.0)
134.4 (130,0-141.0)
132.5 (126,0-137.0)
132.8 (128.0-139.0)
131.6 (125.0-137.0)
136.7 (129.0-142.0)
137.5 (130.0-146.0)
140.7 (135.0-148.0)
138.2 (131.0-148.0)
137.7 (130.0-146.0)
154.0 (151.0-158.0)
136.0 (123,0-143.0)
Culmen
26.3 (24.0-28.0)
25.5 (24.0-27.0)
25.7 (25.0-27.0)
25.9 (24,0-27.5)
25.8 (25.0-27.0)
25.8 (23.0-27.0)
25.4 (24.0-27.0)
25.4 (24.0-26.5)
25.1 (24.0-27.0)
25.6 (24.5-27.0)
32.8 (31.5-34.0)
26.6 (26.0-27.8)
7 MEASUREMENTS IN MILLIMETERS (AVERAGE, MINIMUM AND MAXIMUM) OF THE Paciric Coast Forms or APHELOCOMA
Depth of bill
at nostril
9.2 (8.8- 9.8)
8.2 (8.0- 8.5)
8.3 (8.2- 8.5)
8.8 (8.0- 9.2)
9.1 (8.0-10.0)
9.1 (8.5- 9.5)
9.1 (8.5- 9.2)
9.3 (8.8-10.0)
9.1 (8.5- 9.5)
9.7 (9.2-10.0)
10.1 (10.0-10.2)
8.1 (8.0- 8.5)
Tarsus
38.4 (37.0-40.5)
39.7 (38.5-40.5)
38.6 (38.0-39.5)
39.7 (38.0-41.0)
40.3. (38.0-43,0)
41.5 (39.0-43.0)
40.2 (39.0-41.5)
40.8 (38.0-42.0)
40.7 (39.0-42.0)
41.6 (40.0-44.0)
46.3 (45.5-47.0)
39.5 (38.0-42.0)
Middle toe with-
out claw
23.0 (22.0-23.5)
21.8 (20.0-23,0)
21.8 (21.0-22.5)
22.5 (21.0-23.5)
23.5 (22.0-25.0)
23.8 (22.5-24.5)
24.4 (23.0-26.0)
23.2 (21,0-24.0)
24.0 (23,0-25.0)
24.2 (23.0-25.0)
24.9 (24,0-26.0)
22.1 (21.0-23,0)
422
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UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
: ZOOLOGY
Vol. 17, No. 14, pp. 423-430
* Vol. 17, No. 15, pp. 431-433
April 25, 1918
Me. ie
i hes
14. SIX NEW MAMMALS FROM THE, i
MOHAVE DESERT AND INYO >.”
REGIONS OF CALIFORNIA
BY
JOSEPH GRINNELL
ALASKA AND BRITISH
COLUMBIA
BY
HILDA WOOD GRINNELL
UNIVERSITY OF CALIFORNIA PRESS
BERKELEY
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EE
UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 14, pp. 423-430 April 25, 1918
SIX NEW MAMMALS FROM THE MOHAVE
DESERT AND INYO REGIONS OF
CALIFORNIA
BY
JOSEPH GRINNELL
(Contribution from the Museum of Vertebrate Zoology of the University of California)
Field work carried on under the auspices of the California Museum
of Vertebrate Zoology during 1917 in southeastern California brought
to light many new facts in regard to the general distribution and
speciation of the endemic vertebrate animals. Some of these facts, as
concerning certain of the mammals, are set forth in the present paper.
Scapanus latimanus monoensis, new subspecies
Mono Mole
Type—Female adult, skull (with teeth worn) and skin (in summer
pelage) ; no. 25834, Mus. Vert. Zool.; Taylor Ranch, two miles south of
Benton Station, Mono County, California; August 29, 1917; collected
by H. G. White; original no. 1376.
Diagnosis—A small-sized member of the Scapanus latimanus group
of moles (see Jackson, 1915, pp. 64-75) ; similar to its near neighbor
on the south, S. 1. grinnelli, but color mouse gray (of Ridgway, 1912,
pl. 51) instead of fuseous-black, and size slightly less. Resembles
S. Ll. dilatus in color but size very much less.
Measurements.—Average of eight adults (the first three features as
taken by the collector in the field) : total length, 150 millimeters; tail
vertebrae, 33; hind foot, 20.6; greatest length of skull, 32.9; mastoid
breadth, 16.2; interorbital breadth, 7.5. Extremes are shown in the
accompanying table (1).
Distribution—Known only from two localities, both in Mono
County, California: vicinity of Williams Butte. near Mono Lake; and
vicinity of Benton. The total available material representing this form
is listed in the accompanying table.
[ Vou. 17
University of California Publications in Zoology
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1918] Grinnell: New Mammals from the Mohave and Inyo Regions 425
Thomomys melanotis, new species
White Mountains Pocket Gopher
Type.—Male adult, skull and skin (in nearly full new pelage) ;
no. 26499, Mus. Vert. Zool.; 10,500 feet altitude on Big Prospector
Meadow, White Mountains, Mono County, California; July 27, 1917;
collected by A. C. Shelton; original no. 3402.
Diagnosis—A Thomomys with characters of the ‘‘perpallidus
group’’ in part and of the ‘‘alpinus group’’ in part (see Bailey, 1915,
pp. 33, 63, 68). General coloration very pale, almost identical with
that in 7. perpallidus perpallidus, differing from that form only in
more dusky nose and mouth and in more extensive, slaty black, ear-
patch. Ear extremely small, rounded, and densely clothed with fine
black hairs. General size medium, about as in 7. alpinus alpinus.
Skull relatively narrow, and with slender rostrum and unaccentuated
ridges, thus resembling alpinus; but dentition lighter, interorbital
constriction narrower, and auditory bullae much larger.
Comparisons.—In coloration, much paler and grayer than any of
its geographical neighbors (Thomomys perpallidus perpes, T. scapterus,
T. operarius, and T. alpinus alpinus). Differs from perpes otherwise
in smaller and more hairy ears, extensively black ear-patch, lighter
skull, narrower interorbital constriction, narrower rostrum, slenderer
zygomata, much smaller teeth, and larger auditory bullae. Differs
from scapterus in larger size, extensively black ear-patch, more widely
and more squarely spreading zygomatic arches, broader brain-ease,
much longer nasals, and larger auditory bullae. Differs from operarius
in conspicuously black-haired ears, extensive black ear-patch, lighter
and slenderer skull, much narrower rostrum, more projecting incisors,
much lighter dentition, constricted nasals, narrower interorbital con-
striction, and more inflated auditory bullae. Differs from alpinus in
more extensive black ear-patch, less projecting incisors, much smaller
molar teeth, narrower interorbital constriction, and much larger
auditory bullae.
Measurements.—See Table 2; compare with those of various gophers
as given in Bailey (1915).
Distribution —The nine specimens secured (see table) were all
taken in the Canadian and Hudsonian life-zones, 10,000 to 10,500 feet
altitude, on, and within three miles of, Big Prospector Meadow, White
Mountains, Mono County, California.
Remarks.—The writer confesses his inability, with the material now
available, to determine satisfactorily the relationships of this new
gopher. It was thought at first to belong unquestionably to the alpinus
group and to constitute a sequestered and far differentiated form
within that series. A few gophers from the vicinity of Benton, Mono
County, and elsewhere in the northern end of Owens Valley, however,
show characters which make it seem possible that melanotis has been
derived from the lowland perpallidus stock through altitudinal in-
vasion. Further acquisition of specimens from the Inyo region will
be needed to clear up the question.
[ Vou. 17
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1918] Grinnell: New Mammals from the Mohave and Inyo Regions 427
Thomomys perpallidus mohavensis, new subspecies
Mohave River Pocket Gopher
Type—Male adult, skull and skin; no. 4639, Mus. Vert. Zool.;
Mohave River bottom, 2700 feet altitude, near Victorville, San Ber-
nardino County, California; December 26, 1904; collected by J.
Grinnell and J. Dixon; original no. 906.
Diagnosis—A Thomomys of the ‘‘perpallidus group’’ (see Bailey,
1915, pp. 33, 68). Resembles 7. perpallidus perpallidus eranially, but
color much darker above, bright cinnamon-buff (of Ridgway, 1912,
pl. 29), and tail shorter; differs from 7. p. perpes (topotypes) in
lighter, more cinnamon tone of coloration dorsally, in slightly larger
size, in greater and squarer spread of zygomatic arches, in more pro-
jecting incisors, and in much larger auditory bullae.
Measurements.—See Table 3.
Distribution —Abundant along the bottomlands of the Mohave
River, in San Bernardino County, at least from Victorville down
(north) to Barstow; also, transversely, along the southern rim of the
Mohave Desert, from Cushenbury Springs, San Bernardino County,
west at least to Fairmont, Los Angeles County.
Remarks—As already stated, by Bailey (1915, p. 69), pocket
gophers are not continuously distributed over the desert areas of the
Southwest. Wide areas are unrepresented at all, these animals thus
occurring in more or less remotely isolated colonies, usually where
permanent moisture produces a continuous growth of edible plants.
The larger valleys, like those of the Mohave and Owens rivers, well
separated from one another by desert reaches, have evidently served
as effective differentiation centers, and we find several distinguishable
races of the pocket gopher accordingly. It seems equally true that
some of the higher and more isolated of the mountain ridges of the
TABLE 3
MEASUREMENTS, IN MILLIMETERS, OF EIGHT ADULT SPECIMENS OF Thomomys per-
pallidus mohavensis, ALL COLLECTED BY J. GRINNELL AND J. DIXON IN THE
VICINITY OF VICTORVILLE, SAN BERNARDINO COUNTY, CALIFORNIA
: a & 2 i,
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ies ee) ae ee ee Bea eee
eee Sex Date = = is ge Be a7 58 a ieee Ba
4639" of Dee. 26, 1904 235 75 Bal 36.4 alfay-45) 26.1 20.7 6.4 8.0
4646 dé Dee. 29, 1904 227 76 30) 33.0 12.2 23.0 19.5 6.6 7.8
4660 3 Dee. 31, 1904 223 75 28 34.9 12.9 24.4 20.1 6.2 7.6
4661 fof Dee. 28, 1904 228 66 30 37.0 ye 27.8 21.9 6.7 8.1
4644 2 Dee. 31, 1904 195 62 28 30.6 10.0 22.3 18.1 6.5 7.6
4647 2 Jan, 1,1905 212 60 29 32.3 13.0 23.5 19.8 6.9 Tei
4648 io) Dee. 27, 1904 230 69 30 2.2 12.4 23.4 20.1 6.5 8.0
4650 g Dee. 27, 1904 210 70 29 31.9 13.0 23.7 20.0 6.7 Ti
1 Type.
428 University of California Publications in Zoology | Vou. 17
same general region have operated in similar fashion, because of the
soil-moisture maintained thereon. Thomomys perpallidus mohavensis
and T. p. perpes are valley forms; T. scapterus and T. melanotis are
montane forms.
Perodipus mohavensis, new species
Mohave Kangaroo Rat
Type.—Male adult, skin and skull; no. 26835, Mus. Vert. Zool. ;
3275 feet altitude, half mile east of railway station of Warren (about
five miles north of Mohave), Kern County, California; March 27,
1917; collected by J. Grinnell; original no. 3942.
Diagnosis—A medium sized rather small eared, buff-colored
Perodipus, perhaps nearest like P. panamintinus. Differs from topo-
types of that form in somewhat smaller size, decidedly smaller ears,
much more ochraceous-buffy tone of coloration, in less amount of
black about the face, slightly narrower skull, and in less inflated
auditory bullae.
Measurements.—See table 4 (compare with measurements given by
Merriam, 1894, 1904, and 1907, for the different species of Perodipus
described by him).
Distribution—Specimens at hand indicate the presence of this
form along the western border of the Mohave Desert at least from
Walker Pass, northeastern Kern County, south to Fairmont, north-
western Los Angeles County.
TABLE 4
MEASUREMENTS, IN MILLIMETERS, OF TEN ADULT SPECIMENS oF Perodipus
mohavensis, ALL COLLECTED BY J. GRINNELL AND J. DIXON NEAR MOHAVE,
KERN COUNTY, CALIFORNIA, MARCH 26 AND 27, 1917
= Eo a
= > oe ie ee go Vig siHie
Museum = Pa i He es 2 3 fs = =
number Sex & a q Sine i AQ o =
26835* ref Bihy alyAsh | 2 al 40:7 25.0 15.8 5.2
26837 J 295 175 44 12 39:4 240 “Wb 5:0
26839 Jb 290 175 44 i383 39.2 245 14.6 4.7
26841 Jd 300 180 41 13 40.00 244 15.5 5.3
26843 Jb 305) 180)” 455) 3 4170) 20.38 Lo 8ieSro
26830 2 288) 272) 40) W0l5 (3821 Peay MES) hy
26832 ie) 285 167 43° 12 39.4 248 15.0 4.8
26834 ie) 295) Lid) 425 eel 39.6 240 148 49
26840 2 285 170 43 £14 38.3 23.2 146 4.8
26842 2 300 175 44 138 39.3 24.0 14:9 5.2
1 Type.
1918] Grinnell: New Mammals from the Mohave and Inyo Regions 429
Callospermophilus chrysodeirus perpallidus, new subspecies
Inyo Golden-mantled Ground Squirrel
Type.—Mrale adult, skull and skin (in partially new winter pelage,
otherwise worn breeding pelage) ; no. 27488, Mus. Vert. Zool.; 10,300
feet altitude, near Big Prospector Meadow, White Mountains, Mono
County, California; July 26, 1917; collected by J. Grinnell; original
no. 4334.
Diagnosis.—Resembles Callospermophilus chrysodeirus chrysodeirus
of the Sierra Nevada, but general coloration paler; middle of back,
rump and sides, more ashy in tone, head less richly tawny, and under
surface of body whiter. As a result, the black dorsal stripes give an
impression of greater sharpness. Resembles C. trepidus of the Pine
Forest Mountains, northern Nevada, but tail shorter and coloration
even paler.
Material—F orty-five specimens, from the Inyo and White moun-
tains, 7,000 to 11,600 feet altitude, Inyo and Mono counties, California.
Southernmost station, Mazourka Canon, at 7700 feet altitude, in the
Inyo Mountains directly east of Independence.
Measurements of type—Total length, 265 millimeters; tail verte-
brae, 90; hind foot, 39; height of ear from crown, 13.
Remarks.—This is simply a pale desert-range race, probably cut
off but incompletely from its near relative, chrysodeirus, of the Sierra
Nevada.
Ochotona schisticeps sheltoni, new subspecies
White Mountains Cony
Type.—Male adult, skull and skin (showing chiefly newly acquired
winter pelage) ; no. 27560, Mus. Vert. Zool.; 11,000 feet altitude, near
Big Prospector Meadow, White Mountains, Mono County, California ;
July 29, 1917; collected by A. C. Shelton; original no. 3414.
Diagnosis—Nearest like Ochotona schisticeps schisticeps in general
coloration; tones of color fully as dark dorsally, but belly and tops
of feet less pervaded with tawny; ears blackish, margined more con-
spicuously with white. Cranium as in O. s. schisticeps, O. s. mwiri,
and O. s. albatus (between which three Sierran races there appear to
be no eranial differences) but with notably larger auditory bullae, and
with brain-case higher, more curved dorsally as seen in profile.
Measurements of type—Head and body, 188 millimeters; tail
vertebrae, 8; hind foot, 30; height of ear from crown (inner base), 24;
total weight, 132.5 grams; occipitonasal length of skull, 42.8 milli-
meters; height of brain-case at bullae, 15.6; greatest cranial width
ineluding bullae, 21.8; greatest diameter of bulla (diagonally antero-
posteriorly), 13.7.
Material—Thirty-nine specimens, all taken in the White Moun-
tains, in Mono and Inyo counties, California, at altitudes ranging from
8,200 to 11,900 feet.
430 University of California Publications in Zoology (Vou. 17
Remarks.—Because of the aridity of the White Mountains I had
expected to find the cony on this range relatively light colored, pos-
sibly identical in this regard with the pale-colored race albatus of the
Mount Whitney region. It turns out, however, that the White Moun-
tains animal is darker even than muiri of the Yosemite region, closely
resembling in color tone the race schisticeps of the northern Sierras
from the Tahoe region northward.
As elsewhere, the conies in the White Mountains live in rock slides
and broken-up rock outerops. Two colors of rocks occur in this range,
a blackish or dark red “‘shale,’’? and a white or grayish white granite.
There are extensive belts purely of one or the other kind of rock. Our
party took pains to shoot conies from each color of ground, keeping
notebook record of where each specimen was shot. I am unable to detect
any difference in color between animals shot from white granite and
those from dark ‘‘shale.’’
The new subspecifie name, sheltoni, is selected in recognition of the
efficient services as field collector rendered the Museum of Vertebrate
Zoology by Mr. Alfred C. Shelton. It was chiefly through his tireless
effort that the excellent series of White Mountains conies was obtained.
LITERATURE CITED
BAILEY, V.
1915. Revision of the pocket gophers of the Genus Thomomys. U.S. Dept.
Agric., Bur. Biol. Surv., N. Amer. Fauna, 39, 136 pp., 8 pls.,
10 figs. in text.
JACKSON, H. H. T.
1915. A review of the American moles. Jbid., 38, 100 pp., 6 pls., 27 figs. in
text.
MERRIAM, OC. H.
1894, Preliminary deseriptions of eleven new kangaroo rats of the genera
Dipodomys and Perodipus. Proce. Biol. Soe. Wash., 9, 109-116.
1904. New and little known kangaroo rats of the Genus Perodipus. Ibid.,
17, 139-146.
1907. Deseriptions of ten new kangaroo rats. Ibid., 20, 75-80.
Ripeway, R.
1912. Color standards and color nomenclature. (Washington, published by
the author), 8 + 44 pp., 53 col. pls.
Transmitted March 5, 1918.
UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 15, pp. 431-433 April 25, 1918
NOTES ON SOME BATS FROM ALASKA AND
BRITISH COLUMBIA
BY
HILDA WOOD GRINNELL
(Contribution from the Museum of Vertebrate Zoology of the University of California)
The following notes seem worthy of publication as being contribu-
tory to the very meager knowledge so far available in regard to the
bats of Alaska and British Columbia. Specimens from these regions
are very slow to accumulate; as they do become more plentiful it is
inevitable that previous views in regard to identity and relationships
of the species represented be more or less modified.
Myotis longicrus longicrus (True)
Among the twelve specimens of bats listed from Admiralty and
Baranof islands, Alaska, by Heller (1909, p. 264) under the name
Myotis lucifugus alascensis, there is one which is clearly referable to
the longicrus group. This specimen (4, no. 186, Mus. Vert. Zool.) was
collected by C. Littlejohn at Mole Harbor, Admiralty Island, June 9,
1907. It measures in millimeters: total length, 90.0; tail, 40.0; fore-
arm (both defective) ; tibia, 18.0; foot, 9.0; greatest length of cranium,
14.3; breadth of brain-ease, 7.5. In color this example is deep Vandyke
brown both above and below, the hairs with tippings of cimnamon
brown. The coloration of this specimen is almost identical with that
of examples of longicrus taken in northwestern California. The
characters of this individual throughout thus align it with Myotis
longicrus longicrus.
In his account of a collection of birds and mammals from Van-
couver Island, British Columbia, Swarth (1912, pp. 109, 110) men-
432 University of Califorma Publications in Zoology [| Vou. 17
tions a bat (9, no. 12588, Mus. Vert. Zool.) taken at Errington,
August 31, 1910, which he refers to the form Myotis lucifugus
alascensis, stating that ‘‘the specimen is so imperfect as not to admit
of exact identification.’’ While the tibiae of this bat are mutilated,
it is otherwise in good condition and the skull is perfect, showing the
up-turned rostrum and the facial angle typical of Myotis longicrus
longicrus. The forearm of this bat measures 36 millimeters, the
greatest length of cranium, 14.3, and breadth of brain-case, 7.5. In
color this example is very close to the specimen from Admiralty
Island, Alaska, described above, but the hair tippings both above and
below are lighter, more ochraeous-buff.
In so far as the writer is aware Myotis longicrus longicrus has not
hitherto been recorded from farther north than the type locality,
Puget Sound, Washington.
Myotis lucifugus alascensis Miller
Among some bats belonging to the Biological Survey collection,
United States National Museum, and loaned to the writer through the
courtesy of Mr. E. W. Nelson, are four skins (three with skulls) from
Skidegate, Queen Charlotte Islands, British Columbia. These speci-
mens (nos. 100675-100678) are adult males collected by W. H. Osgood
in July, 1900, and are presumably the four recorded (Osgood, 1901,
pp. 36, 37) under the name Myotis ywmanensis saturatus. The three
skulls measure in millimeters: no. 100675, greatest length of cranium,
14.7; breadth of brain-case, 7.3; no. 100676, greatest length of cranium,
14.7; breadth of brain-case, 7.6; no. 100677, greatest length of cranium,
14.9; breadth of brain-case, 7.4. The specimens in all their characters
show their identity with the bat now currently known as Myotis
lucifugus alascensis.
Myotis californicus caurinus Miller
There are in the collection of the Museum of Vertebrate Zoology
two mummified specimens of Myotis californicus caurinus collected by
W. D. McLeod at Howkan, Long Island (near Dall Island), south-
eastern Alaska. These bats measure in millimeters: no. 19292 (skull
removed) : forearm, 31.7; tibia, 12.3; thumb, 3.6; greatest length of
skull, 13.1; breadth of brain-case, 6.8. No. 19293: forearm, 32.2;
tibia, 13.5; thumb, 3.9. In color these two examples are closely
1918] Grinnell: Some Bats from Alaska and British Columbia 433
similar; the fur is everywhere plumbeous-black at the bases of the
hairs and deep Prout’s brown at the tips; under surface slightly lighter
than dorsal surface.
This appears to constitute the first record of Myotis californicus
caurinus from Alaska.
LITERATURE CITED
HELLER, FE.
1909. The mammals in Birds and mammals of the 1907 Alexander Expedition
to southeastern Alaska. Univ. Calif. Publ. Zool., 5, 245-264, pl. 26,
text figs. 3, 4.
Oscoop, W. H.
1901. Natural history of the Queen Charlotte Islands. U. 8. Dept. Agric.,
Div. Biol. Surv., N. Amer. Fauna, 21, 7-50, pls. 1-5, text fig. 1.
SwartH, H. 8.
1912. Report on a collection of birds and mammals from Vancouver Island.
Univ. Calif. Publ. Zool., 10, 1-124, pls. 1-4.
Transmitted March 5, 1918.
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ZOOLOGY
Vol. 17, No. 16, pp. 435-504, plates 25-29, 16 text figures May 29, 1918
REVISION OF THE RODENT GENUS
APLODONTIA
BY
WALTER P. TAYLOR
(Contribution from the Museum of Vertebrate Zoology of the University of California)
CONTENTS
PAGE
Ne UIE ROY6 ELOISE Re Ee ea 435
1. Prefatory remarks .. 435
2. Material and acknowledgements so HIBY/
3. Methods of measurement . 438
SMe Vise VOM TPAD LOM OTULIG) ox oreo oe eerecee eens eee eee . 439
1. Age variation 439
PR SO MUA OUITTCTONCOS | aercs cee eens sansa scan feceeewsarcwensiecscacasertds suevereese?screveresce 40
3. Individual variation 441
+. Molt and seasonal variation 445
Si, GuexOpSe OLOGY ON ace cater ree eee ec eet cee o are a EO 445
(CL Tabsysyitomeaye One TN Nj ol kay Moya ASS eee ce eae en . 447
Tl. Gaerne) HOMES) cee eer eres eo RE 447
Pea SS LICE TNE A revs AE At LS CU SSO WN ess ae ee renee an eee eee tee
TD}, TSUPA RS; OIE a UNG P OTA TOT es oe see ee EE ne eee eee er ee
E. Present systematic status of Aplodontia
Tle, Mblanes sigemeawillies PAN oy Kaye (oars 0G eRe oc ee 4
2s VIB ave Sera SS Za GYE NO HI OT, ee eccE e esc cereeCERES ° 452
3. Variations in the generic name -_.. 52
4. List of species and subspecies with type localities -......-.-.--.--.. 453
Sis LNCS) OIE SOLEMN enceee ecco ere ee ce aE ea Sane encore 454
EF.
Literature cited, with titles of other works containing matter on Aplodontia 484
A. INTRODUCTION
1. Prerarory REMARKS
Discovered by Lewis and Clark in the early years of the nineteenth
century, the peculiar west American rodent genus Aplodontia has
long attracted interest and attention. It is the sole living representa-
tive of that primitive group of rodents, the Aplodontoidea (see
Matthew, in Osborn, 1910, pp. 534, 535), from which all other mem-
436 University of California Publications in Zoology — (Vou. 17
bers of the order have been presumed to be derived, and constitutes
a stock which is one of the most conservative known in the class
Mammalia. The family to which it belongs (the Aplodontiidae) was
the sole mammalian family cited by Wallace (1876, p. 127), as char-
acteristic of his Californian subregion of Nearctica, and is remark-
able for its present restricted distribution, being found only on the
Pacific coast of North America, in an area which may be bounded
roughly by southern British Columbia on the north, the Sierra-
Cascade mountain system on the east, and the latitude of San Fran-
cisco Bay on the south (see map, text-fig. H).
The unusual characters of the genus Aplodontia (see pls. 26, 27),
together with the scarcity of fossil remains representative of it, have
made the determination of its systematic status a matter of exceptional
difficulty. Minding (1829, p. 86) referred the genus Anisonix
[= Anisonyx = Aplodontia (part)| to the family Prensiculantia,
in which it is associated with twenty-seven other genera, ranging from
Castor, Hydromys and Mus to Bathyergus, Tamias and Chiromys.
Two years later Bonaparte (1831, p. 20) associated the genus Aplo-
““section’’ Sciurina of the
family Muridae. Swainson (1835, p. 388) put Aplodontia in his
“Division I’’ of the Order Glres which contained twenty-two addi-
tional genera. By Schinz (1845, pp. 120, 139) the genus was referred
dontia with twenty other genera in his
to the family Cunicularia, in which were included also arvicolines,
octodonts, pocket gophers, and mole-rats. Gervais (1854, p. 364)
grouped Aplodontia with the pocket gophers. Baird (1857, p. 350)
included Aplodontia in the subfamily Castorinae; Lilljeborg (1866,
opp. p. 9) referred it to the porcupines; Peters (1865, pp. 177, 179),
Alston (1876, p. 66, and pl. 4, opp. p. 61), Coues (1877), pp. 543,
601) and others placed it with the squirrels. Guill (1872, p. 22) refer-
red the Haploodontidae to a superfamily Haploodontoidea, listing it
as equivalent in rank to the Castoroidea and Sciuroidea ; Zittell (1894,
p. 528) included the Haplodontidae with certain aberrant families of
rodents in his Protrogomorpha; and Thomas (1896, pp. 1014, 1015)
accorded to the Aplodontiidae and Anomaluridae separate group
rank, leaving it for further research to show their true relationships.
The latest classification is that of Matthew and Osborn in 1910,
according to which the Aplodontiidae are associated with three extinct
families of rodents in the superfamily Aplodontoidea.
Thirteen forms of Aplodontia Recent have been deseribed, nine
being here recognized. The first formal description was that of
1918] Taylor: Revision of the Rodent Genus Aplodontia 437
Rafinesque, who characterized ‘‘ Anisonyx? rufa’’ in 1817 (p. 45) on
the basis of Lewis and Clark’s account of the sewellel as observed in
the neighborhood of the Columbia River. In 1829 Richardson (1829a,
pp. 333-837) deseribed Aplodontia leporina on the basis of specimens
collected by David Douglas from within the range of what is now
regarded as Aplodontia rufa rufa. For thirty-six years no further
new names appeared. In 1865 (pp. 177-179) Peters named Haplodon
leporinus var. Californicus from a specimen received at the Berlin
Museum, said to have come ‘‘aus den Gebirgen Californien.’’ It is the
opinion of the writer that this name is tenable for the aplodontia of
the Sierra Nevada Mountains of California, which was described by
Dr. C. Hart Merriam in 1886 (p. 316) under the name Aplodontia
major. Four forms of Aplodontia were deseribed by Dr. Merriam
in 1899a (pp. 19-21), as follows: Aplodontia pacifica, type locality,
Newport, Oregon; Aplodontia phaca, type locality, Point Reyes, Cali-
fornia; Aplodontia olympica, type locality, Quiniault Lake, Washing-
ton; and Aplodontia major rainiert, type locality, Paradise Creek,
Mount Rainier, Washington.
In 1914 two additional forms were described from California:
Aplodontia chryseola (here referred to rufa), from Jackson Lake,
Siskiyou County, by Kellogg (p. 295), and Aplodontia mgra, from
Point Arena, by the present writer (p. 297). Two years later three
more forms were characterized by the writer: Aplodontia californica
columbiana, from Roab’s Ranch, near Hope, British Columbia (Tay-
lor, 1916c, p. 499) ; Aplodontia rufa grisea (1916c, p. 497, here refer-
red to rufa), from the vicinity of Seattle, Washington; and Aplo-
dontia humboldtiana (1916a, pp. 21-24), from the Humboldt Bay
region, California.
2. MATERIAL AND ACKNOWLEDGMENTS
The work here reported upon was begun while the writer was a
staff-member at the Museum of Vertebrate Zoology, University of
California, and finished after his appointment to the staff of the
Bureau of Biological Survey, Washington, D. C. Exclusive of fossil
material the present study is based on the examination of 369 speci-
mens, for the most part skins with skulls contained in the collections
of the two institutions named.
Through the courtesy of Professor John C. Merriam the writer
has had access to the University of California Collections in Vertebrate
438 University of California Publications in Zoology — | Vou. 17
Palaeontology, in which are contained some of the most interesting
and important specimens known bearing upon the history of the
Aplodontiidae and the apparently related fossil family Mylagaulidae.
Acknowledgments are due the following persons: For the loan of
additional material for study, to Messrs. F. J. V. Skiff and Wilfred
H. Osgood, of the Field Museum of Natural History, Dr. John F.
Bovard and Mr. Alfred C. Shelton, of the University of Oregon
Museum, and Messrs. Samuel Henshaw and Outram Bangs, of the
Museum of Comparative Zoology; for the courtesy of access to the
collection under his charge, to Mr. Gerrit S. Miller, Jr., of the United
States National Museum; for helpful suggestions regarding the his-
torical problems involved, to Dr. John C. Merriam, of the University
of California; for various suggestions in connection with the work,
to Messrs. E. W. Nelson, T. S. Palmer, Vernon Bailey, Edward A.
Preble, and Hartley H. T. Jackson, of the Biological Survey, and to
Messrs. Gerrit S. Miller, Jr., and J. W. Gidley, of the United States
National Museum; for a list of vernacular names of Aplodontia used
by California Indians, to Dr. C. Hart Merriam, of the Harriman
Foundation, Smithsonian Institution; and for eritical oversight, to
Dr. Joseph Grinnell, of the Museum of Vertebrate Zoology, University
of California, at whose instance the work was undertaken.
3. MretrHops or MEASUREMENT
All measurements are in millimeters.
Cranial measurements, except where otherwise specified, are taken
as follows:
Basilar length: inferior lp of foramen magnum to posterior
margin alveolus of incisor.
Length of nasals: most anterior point on nasal bones to most
posterior point.
Width of nasals: greatest distance across both of them.
Length of audital tube: outer border of foramen ovale to farthest
lateral point (with reference to the skull) on zygomatic (or anterior)
side of audital tube.
Length of incisive foramina: greatest length of foramen on right
side, that is, with skull resting on its dorsal surface and rostrum point-
ing away from the observer.
Zygomatic width: greatest inclusive measurement, taken outside
of the zygomatic arches.
1918] Taylor: Revision of the Rodent Genus Aplodontia 439
Greatest width of interpterygoid fossa: at expansion of fossa
immediately back of hard palate.
Mastoid width of cranium: greatest inclusive measurement taken
outside of mastoid processes.
Alveolar length superior cheek teeth: most anterior point on
alveolus of premolar four to most posterior point on alveolus of molar
three.
Distance between infraorbital foramina: measured on ventral
surface of skull.
Mandible, transversely across angular process: greatest dimension
along axis of process, nearly at right angles to axis of mandible itself.
Greatest length of mandible: most posterior point on articular
condyle of mandible to most anterior point on alveolus of incisor.
External measurements are ordinarily taken as given below. The
short and well-haired tail of Aplodontia makes the determination of
the total length a matter of some difficulty, in consequence of which
this measurement has sometimes been taken on the skinned body
instead of as specified below.
Total length, on unskinned body stretched out lengthwise, most
anterior point on cartilage of nose to tip of tail, exclusive of hairs.
Hind foot, heel to tip of longest claw.
B. VARIATION IN APLODONTIA
. 1. Ack VARIATION
As the aplodontia grow older the soft gray pelage of the young
animal becomes less soft and more brownish or blackish, a ventral
brown wash may appear in the adult though seldom in evidence in
the young, and the numerous white-tipped hairs which stand out so
conspicuously in the pelage of the juvenal become obscured. That
specific differentiation takes place early is indicated in several of
the subspecies, in none more strikingly, however, than in Aplodontia
rufa nigra, in which the black coloration of the adult is noted in
animals of the vear taken in July.
The following tendencies may be observed in the crania as maturity
approaches. In dorsal view (pl. 25) all the sutures but those bound-
ing the nasal bones laterally tend to disappear; the interorbital con-
striction tends to grow narrower, proportionally and absolutely ; the
temporal lines or ridges become accentuated and approach one another,
440) University of California Publications in Zoology — (Vou. 17
though never, apparently, forming a true sagittal crest (the degree
of their approach is different in different species) ; the mastoid pro-
cesses tend to grow laterad more rapidly than do the audital tubes;
the lambdoidal ridge undergoes marked development; the skull
becomes more flat, less round, changing from the more squirrel-like
form of early youth to the more specialized Aplodontia type of full
maturity ; all processes become accentuated, and the angular process
of the mandible undergoes a considerable transverse expansion, its
development proceeding at such a rate that the width of the mandible,
measured along the axis of this process, increases faster proportion-
ally than does the length of the mandible. Through all these changes
the distance anteriorly across the palate between the alveoli of the
fourth premolars remains practically constant.
The permanent teeth are long-crowned and as soon as they become
somewhat worn afford no elue to age. The tooth formula of Aplo-
=o las ONOM ARO ens
dontia is 1 oS an Bae 3
specimens is as follows: Superior, milk P*, milk P*, M’, M*, M*,
Ore Tooth eruption in available
permanent P*, permanent P*; inferior, milk P,, M,, M., M., per-
manent P,.
The fourth premolars (P*, P,), are somewhat less specialized in
the deciduous dentition than in the permanent. They are brachyo-
dont, somewhat tubereulated, and have deep enamel lakes, in the
former; hypsodont, with tubereulation obscure, and with shallower
enamel lakes, all trace of which is soon lost by wear, in the latter.
The considerable variation in size of cranium, as well as in the
weight of its bars and processes, noticeable in series of fully adult
skulls from the same general locality, indicates that slow growth may
continue throughout life.
2. SexusAL DIFFERENCES
Aside from a not well-marked tendency toward larger measure-
ments on the part of the males (which, in specimens of Aplodontia
rufa phaca measured, average eight per cent longer than the females),
and the presence of a series of conspicuous ventral markings about
the mammae of summer females, practically no differences due to
sex can be made out externally. There are in this genus three pairs
of mammae. About each, in females in summer pelage, is a nearly
circular avea of black or dark brown hair ten to twenty millimeters
1918 | Taylor: Revision of the Rodent Genus Aplodontia 44]
in diameter, which stands out in strong contrast to the grayish or
weakly brownish coloration of the underparts. In males and in
ce
winter females the ‘‘spot marks’’ are inconspicuous or lacking.
It would be difficult, if not impossible, to determine the sex of the
individual in any given instance by study of the cranium. There is
a tendency for ridges and processes to be somewhat more accentuated
in the males, for zygomatic arches to be somewhat heavier, and for
the temporal ridges to be more closely approximated. Usually the
males have basilar length, zygomatic width, mastoid width, and dis-
tance transversely across the angular process of the mandible greater
than in the females. The largest, heaviest specimens in any adequate
series of skulls are usually those of males. In Aplodontia rufa
olympica the range of variation in size is greater, on the basis of the
specimens measured, in the males than in the females. In certain
forms the females have the alveolar length of the superior cheek teeth
greater than in the males, though the measurements of the latter may
exceed those of the former in most respects. In at least two forms,
on the basis of measurements taken, the females have interpterygoid
fossae averaging broader than in the males. In practically all
instances, however, the range of individual variation is so great as
to transeend that due to sex.
3. INDIVIDUAL VARIATION
While in the present study it has been impossible to eliminate
geographic, sex, and age variation altogether, still it is believed that
the observations, measurements, and percentages given are of value,
sinee they suggest the range of variation, chiefly individual, which
must be reckoned with in using a typical series of adults in specifie
comparisons.
In a series of eleven specimens of Aplodontia rufa pacifica taken
in February, March and April, there is but little variation in color.
Dorsally and laterally all are grizzled pinkish cinnamon, with the
brown coloration a little more intense in certain specimens, a little
less so in others. The brown wash ventrally varies from near hight
pinkish cinnamon to pinkish buff. A little more variation, however, is
apparent in twelve examples taken during October, November and
December. The dorsal coloration in these varies from cinnamon or
sayal brown to pinkish cinnamon or light pinkish cinnamon. The
pelage of one specimen (no. 9077, Field Mus. Nat. Hist.) is in very
442 University of California Publications in Zoology (Vou. 17
poor condition, suggesting that its aberrant paleness may be due to
disease. As with the early spring examples, the uniformity of this
late fall series of skins is more noteworthy than the variation observed.
Total length and length of tail vertebrae in this genus, as recorded
by the collector, are often unreliable. This follows from the fact that
the condition of the tail, which is very short but well-haired, makes
accurate measurement of tail vertebrae difficult. The writer has,
therefore, put the emphasis in this discussion on the more reliable
cranial measurements.
In rodent species with hypsodont teeth the age of adults is deter-
minable not at all or only with difficulty. In such species reliance
5 10 15 20 25 30 35 40
=
Length of nasals
Width of nasals
Length of incisive foramen
Zygomatie width
Greatest width interpterygoid fossa
Mastoid width
Alveolar length superior cheek teeth
Distance between infraorbital foramina
Mandible, transyersely across angular process
Greatest length of mandible
Fig. A. Diagram showing the percentage range of variation (figured on the
mean) in cranial measurements of twenty-three skulls of Aplodontia rufa pacifica
(see table of measurements page 468).
must be placed on the condition of the sutures, the outline of certain
bones, the form of the skull, and the development of ridges and pro-
cesses. It follows that it is possible that an observed range in varia-
tion assumed to be individual may really be due to age; and this
point ought to be kept in mind through the following discussion.
The diagram (text-fig. A) illustrative of the percentage range of
variation shows that for specific comparisons the basilar length, length
of nasals, width of nasals, mastoid width, and the width of the
mandible, transversely across angular process, are the most depend-
able characters. Zygomatie width and alveolar length superior cheek
1918] Taylor: Revision of the Rodent Genus Aplodontia 443
teeth are subject to greater variation, while the length of the incisive
foramina, greatest width of interpterygoid fossa, and distance between
infraorbital foramina are subject to a considerable range of varia-
tion. It is doubtless not an accident that the measurements which,
in the nature of the case, can be taken with greatest precision, as
basilar length and mastoid width, appear to be subject to less varia-
tion than those which are more difficult to take with entire accuracy.
The outline of the nasals is fairly uniform in the entire series, the
small variation observable being in the condition of the shallow embay-
ment in the lateral outline of the nasals posteriorly, and in their
relative breadth anteriorly and posteriorly. The width of the inter-
Figs. B-D. Posterior view crania of Aplodontia rufa pacifica, to illustrate
variation in relation of parts. X 1. Fig. B, no. 89309, Biol. Surv. Coll.; fig. C,
no. 9052, Field Mus. Nat. Hist.; fig. D, no. 77371, Biol. Surv. Coll.
orbital constriction is highly variable, not only in animals of different
ages, but also in animals of apparently similar age. In certain speci-
mens a little rounded notch is developed in the lachrymal region, on
the anterior border of the orbit. In others, this notch is feebly
developed or lacking entirely. The degree of development and ap-
proach of the temporal lines or ridges is variable. In some specimens
the ridges are weakly developed and comparatively far apart, in
others strongly developed and much closer together. The weight of
444 University of California Publications in Zoology — (Vou. 17
the zygomatie arch is often different in animals of apparently the
same age. The arches are bowed outward more in some specimens
than in others. The postzygomatie noteh is variable. The degree
of development of the lambdoidal crest varies in animals of apparently
comparable age, being accentuated in certain specimens and setting
off a considerable fossa anterior to it, but in others being compara-
tively weakly developed.
Perhaps the most variable of the cranial characters are furnished
‘by the bones on the posterior aspect of the skull (see text figs. B, C, D).
In the adult, the sutures between the exoccipitals, supraoccipital, and
the basioccipital have completely disappeared. The paroccipital pro-
cess and the oceipital condyle belong to the exoecipitals. The sub-
stance of the mastoid process, as viewed on the posterior aspect of the
cranium, results from the ankylosis of two elements, one, the inner,
from the exoecipital, and the other, the outer, from the squamosal.
A plate from the latter bone invades the outline of the mastoid process
from below, and is marked off by prominent sutures in animals of
every age. The degree of expansion of the winglike mastoids, the
degree of development of the paroccipital processes, and particularly
the degree of invasion of the plate of the squamosal which comes up
from below, are subject to variation, the wide range of which suggests
that it may be due to age. Individual differences are observable in
the outhne of the foramen magnum, which is rounder in some speci-
mens, flatter in others. In the ventral aspect of the skull one notes
that there is considerable variation in the diameter of the third upper
premolar, and noticeable variation in the outline of the hamular pro-
cesses, the length of the incisive foramina, the outline of the inter-
pterygoid fossa, the outlne of the zygomatic arch particularly poste-
riorly, the caliber of the audital tubes and the degree of expansion
and development of the paroccipital processes. The relations of the
foramina in the regions of the foramen lacerum medium are subject
to much variation.
Looking at the skulls in their anterior aspect it is of interest to
note the variation in size of the infraorbital foramen, which in its
ereatest diameter, ranges, in different specimens, from about 2.6 to
4.0 millimeters.
The outline of the coronoid process of the mandible varies from
a form in which it is searcely hooked backward at all, to one in which
the hook form is prominent. The mass and outline of the econdylar
process undergo considerable change from one individual extreme to
1918 | Taylor: Revision of the Rodent Genus Aplodontia 445
the other. The inner and outer prominences of the broad angular
process of the jaw are differently outlined in different specimens,
and the ridge extending from the inner prominence of the angular
process exteriorly on the jaw to bound the masseteri¢ fossa anteriorly
is comparatively well developed in certain specimens, whereas in
others it is interrupted by a smooth space.
4. Motr AND SEASONAL VARIATION
There is but one molt annually in Aplodontia. Pelage renewal
begins in July and August, rarely as early as June, and continues
for two or three months. There is no hard and fast manner of molt-
ing. The hair usually begins coming in on the sides posteriorly and
on the back of the head and neck about the same time. From these
centers the molt spreads until the new pelage covers the body, the
hair of the shoulders and rump being the last to be renewed dorsally.
In some examples the molt proceeds somewhat irregularly. The molt
of the underparts lags behind that of the upperparts. Additional
details as to molt are mentioned in the accounts of species and sub-
species.
The new pelage is longer and sometimes slightly different in color-
ation from the worn pelage it replaces. Thus in Aplodontia rufa
pacifica, the fresh pelage is more richly colored than the worn pelage ;
in A. r. californica, it is a trifle browner; and in A. r. phaea, it is a
little more intensely colored. Differences in coloration and general
appearance are small, however, and in several forms of Aplodontia,
as at present represented in the collections examined, cannot be shown
to exist at all.
5. GmoGRAPHIC VARIATION
Geographic variation, like time variation, is comparatively slight
in the genus Aplodontia. The forms are so little differentiated that
variation geographically is often obscured by individual variation.
All of the described forms may with entire propriety be referred to
a single species.
Mountain forms are larger as a rule than nearby lowland or
coastal forms. Thus Aplodontia rufa columbiana of the mountains
of Hope District, British Columbia, tends to be larger than A. r. rufa
of the Puget Sound region; in lke manner raimiert from Mount
446 University of California Publications in Zoology | Vou. 17
Rainier, Washington, averages larger than rufa; and californica,
inhabiting the Canadian zone of the Sierra Nevada-Caseade moun-
tain system in California, is considerably larger than phaea, found
in the coastal Marin County near San Francisco Bay. Although
this tendeney toward variation in size is slight, its apparently uniform
association with difference in altitude or life-zone indicates its possible
significance.
It has been considered (C. Hart Merriam, 1899a, p. 21; Taylor,
1916c, p. 501) that the ‘‘mountain top’’ subspecies of Aplodontia
(columbiana, rainiert, californica) are more closely related to each
other than to any other forms of the genus. This is possibly true of
columbiana and rainieri (see pl. 28), but the balance of the evidence,
as derived from the present study of rainiert and californica, favors
rather the view that these mountain subspecies are more closely related
to lowland or neighboring forms than to each other. Their general
similarity in size and certain other characters would seem to be due
to parallelism.
Geographic isolation appears to be intimately associated with
speciation in the group. Although material at hand is not sufficient
to demonstrate each step in the process, it is enough to suggest that
the ranges of all the described subspecies of rufa, except nigra and
phaea, mosculate at one point or another. Geographic variation is
continuous, though shght and very gradual, in the intergrading forms.
The most strikingly colored form is the apparently completely isolated
Aplodontia rufa mgra of Point Arena, California.
It is to be noted that whereas on the northern coast of California
three well-marked forms of Aplodontia are found, on a longer coast-
line in the state of Oregon there occurs but one. The three, communi-
ties of Aplodontia on the coast of northern California are separated
from each other by considerable gaps, while along the coast of Oregon
the animals appear to be continuously distributed.
It seems to the writer that the geographical distribution of the
different subspecies of Aplodontia, as well as the degree of develop-
ment and nature of the characters separating them, indicate that
among the factors possibly concerned in their differentiation, geo-
graphic isolation is, at least, one of the most important.
1918] Taylor: Revision of the Rodent Genus Aplodontia 447
C. HISTORY OF THE APLODONTIIDAE
1. GENERAL REMARKS
This family is exclusively North American in origin, development,
and present distribution. In the absence of any other evidence of
European relationship it is probable that the resemblances noted
between the American Oligocene aplodontid genus Allomys and the
genus Sciurodon of the European Oligocene (see Schlosser, 1884, pp.
73, 136) are indicative of accidental convergence rather than real
relationship.
The earliest aplodontid genius is Allomys Marsh (1877, p. 253) or
Meniscomys Cope (1878, p. 5). Found typically in the Oligocene
deposits of the Middle John Day in Oregon, there is only a single
record, so far as known to the writer, of its occurrence elsewhere,
that of Matthew (1904, p. 263), who has recovered an undetermined
species from the Lower Miocene of South Dakota.
The members of the genus Allomys were small, ranging from one-
half to two-thirds the size of Recent Aplodontia. They were much
more squirrel-like than is the Recent genus.
Founded upon a single specimen, an imperfect cranium without
lower jaws, taken in the gravels and tuffs at the top of the Upper
John Day, Mylagaulodon is one of the most interesting of the genera
which are associated with the Aplodontiidae. The characters of the
fourth premolar and infraorbital region of this genus are regarded as
demonstrating its aplodontid position. *
The earliest known fossil remains of the genus Aplodontia were
found by parties from the University of California in the Virgin
Valley Miocene and Thousand Creek Pliocene of northern Humboldt
County, Nevada. The species there recovered is known as Aplodontia
alexandrae Furlong (1910, pp. 397-403). It is somewhat smaller than
the Recent members of the genus, and differs from them in several
particulars, the most important beimg the relative position of the
prominent style internally on the lower molars. For all that, one is
impressed with the resemblances rather than the differences between
the Tertiary Aplodontia alexandrae and the Recent species of the
genus.
Another species of Aplodontia has recently been described by Dr.
J. C. Merriam (1916, pp. 177-179) from the Cedar Mountain region
of western Nevada. The formation in which the specimen (a single
448 University of California Publications in Zoology — |Vou. 17
upper premolar four, somewhat broken) was found has been referred
to the Upper Miocene (loc. cit., p. 171), the Cedar Mountain fauna
being regarded as intervening in time between the Mascall Middle
Miocene of the Middle Basin area and the Barstow Upper Miocene
of the Mohave area.
That both Aplodontia alerandrae and the Cedar Mountain species
represent advanced stages of development between Allomys and
Aplodontia seems clear. Both are nearer Aplodontia than Allomys;
but neither one is so specialized as are the Recent forms of Aplodontia.
There remains to be considered only Aplodontia major fossilis
Sinclair (1905, pp. 147-148), discovered in 1902 by an expedition from
the University of California, in the Pleistocene Potter Creek Cave,
Shasta County, California. Comparison of the numerous remains of
this species in the University of California Collections in Vertebrate
Palaeontology with a large series of skulls and jaws of Aplodontia
rufa californica, the Recent form occurring in the same general
region, shows that the Pleistocene form is very doubtfully if at all
distinguishable from the living species.
2. SUMMARY AND Discussion
It is worthy of remark that nearly all of the records of extinct
members of the family Aplodontiidae are from the now arid territory
east of the great Cascade-Sierra mountain system, which system, at
the present time, marks the eastern boundary of the range of the
genus (see map, text-fig. E). The members of the chiefly Oligocene
genus Allomys hail from the John Day Beds of eastern Oregon, with
one outlying undetermined species in the Rosebud Lower Miocene of
South Dakota; Mylagaulodon comes from the top of the Upper John
Day in eastern Oregon; Aplodontia alerandrae oceurs in the arid
Virgin Valley and Thousand Creek beds of northern Humboldt
County, Nevada; an Aplodontia of an unnamed new species comes
from the Cedar Mountain region of western Nevada; and only Aplo-
dontia major fossilis, found in the Pleistocene Potter Creek and
Samwel caves, in the Shasta region of northern California, comes
from the Pacific slope of the Sierran system. Apparently the range
of the family Aplodontiidae, as well as of the genus Aplodontia, was
formerly much greater than at present, though it must be conceded
that very little is known of the precise relationships of the early
aplodontids to the Recent genus, and we have little data on the west-
ward range of members of the family in Tertiary time.
1918 | Taylor: Revision of the Rodent Genus Aplodontia 449
Reference has already been made to the conservatism of the aplo-
dontid phylum. If, as seems highly probable, Aplodontia alerandrac
is near the direct line of descent of the recent species, it is noteworthy
that, during a lapse of time sufficient for the evolution of the horse
from the Miocene MWerychippus type to the larger Pliocene Pliohippus,
there was no great specific change; and that, during a time which
sufficed to transform the short-toothed, small-sized Merychippus ito
the much larger modern Equus, with modifications in every bone in
the body, and with characters generically probably twice removed, the
aplodontid stock has undergone comparatively slight change, all
Fig. E. Continent of North America, to indicate present range of genus
Aplodontia, and apparent extension of range of family Aplodontiidae in Tertiary
times.
Cross-hatehed area indicates limits of range of Recent genus; cirele represents
record of fossil genera Allomys and Mylagaulodon; triangle indicates an addi-
tional record of Allomys; squares indicate records of genus Aplodontia fossil.
450 University of California Publications in Zoology — (Vou. 17
observed variations, as at present recognized, falling within the limits
of a single genus.
D. HABITS OF APLODONTIA
Aplodontia is herbivorous, colonial, nocturnal, and fossorial. A
considerable degree of humidity and an abundant supply of food
plants seem to be necessary conditions to its existence. Situations well
sheltered by a tangle of vegetation are usually chosen for its burrows.
Its nest is made underground, in an enlarged chamber. Attention
has been called to a hay-making instinct. The “‘hay’’ consists simply
of green plants of various kinds eut up and spread out as if to dry
and to be used later. Sight and hearing are apparently defective;
but smell and touch, particularly the latter, appear to make up for
any deficiencies in these respects.
Little is known of the animal’s breeding activity. Young have
been taken throughout the summer season. It has been asserted that
Aplodontia has a shrill ery. Among its enemies are weasels, skunks
of two genera, wildeats, mink, gray foxes, golden eagles and great
horned owls. Other potential enemies are coons, badgers, and fishers.
Although locally Aplodontia does some damage to man’s interests,
its habitat is such that for the most part it is of no economie signi-
ficance. It burrows holes in ditch walls along the line of the Southern
Pacific in the Sierra Nevada in California; in Oregon it undermines
government trails, causing them to be washed out ; and Lantz (1917, p.
16) reports that in western Washington considerable complaint has
been made of their depredations on crops, particularly small fruits.
The skins are of little or no value.
The habits of Aplodontia have recently been studied by Anthony
(1915, pp. 53-63). The most complete paper on this subject to date
is that of Mr. Charles Camp, now in press in this series of publications.
EK. PRESENT SYSTEMATIC STATUS OF APLODONTIA
1. THe Faminy APLODONTIIDAE
Prensiculantia (part) Minding (1829), p. 86.
Muridae (part), Bonaparte (1831), p. 20.
Sciurina (part), Bonaparte (1831), p. 20.
Cunicularia (part) Wagner (1843), pp. 357, 395.
Pseudostomides (part) Gervais (1854), p. 364.
i =
1918] Taylor: Revision of the Rodent Genus Aplodontia 451
Saccophoriens (part) Gervais (1854), p. 364.
Sciuroides (part) Brandt (1855), p. 151.
Haploodontini or Prismatodontes Brandt (1855), p. 151.
Sciuridae (part), Baird (1857), pp. 240, 350.
Castorinae (part) Baird (1857), p. 350.
Haploodontidae Lilljeborg (1866), table opp. p. 9.
Haploodontoidea Gill (1872), p. 22.
Haplodontidae Alston (1876), pp. 66, 75, 78.
Haplodontiidae Cope (1883), p. 54.
Aplodontidae Allen (1892), p. 31.
Aplodontiae Thomas (1896), p. 1015.
Aplodontiidae Thomas (1896), p. 1015.
Haplodontoidea Weber (1904), p. 496.
Aplodontoidea Matthew, in Osborn (1910), p. 534.
Characterized by Matthew (1910, p. 69) as follows:
1.0.2.3
TONES?
stylids. No postorbital process; zygomatic arch slender; skull wide posteriorly;
postero-inferior portion of angle greatly inflected, posterior end everted. Foss-
orial, Oligocene to recent. Genera, Aplodontia, Meniscomys [= Allomys], Myla-
gaulodon.’’
«Teeth progressively hypsodont, with prominent mesostyles and meta-
Fig. G
Fig. F. Allomys liolophus Cope, dorsal view of cranium. X 1. Univ. Calif.
Coll. Vert. Palae., no. 1672; locality 898, John Day Beds, Oregon.
Fig. G. Allomys cavatus Cope, dorsal view of cranium. X 1. Univ. Calif.
Coll. Vert. Palae., no. 1444; locality 864, John Day Beds, Oregon.
If Allomys is to be referred to the family Aplodontiidae a change
in the above characterization will be necessary ; for postorbital pro-
cesses are present in specimens of Allomys liolophus and A. cavatus
in the University of California Collections in Vertebrate Palaeont-
ology ( see text-figs. F', G).
452 University of California Publications in Zoology — [Vou 17
2. Tur Genus Aplodontia
2
1
3 ; ; ;
we x 222; permanent teeth hyposodont, of secondarily simple
The chief generic characters are as follows: dentition ite+
enamel pattern, characterized by prominent styles internally on the
inferior teeth and externally on the superior teeth; cranium greatly
depressed; postorbital processes on frontal absent; postorbitals on
jugal weakly or not at all developed; infraorbital foramen primitive,
not transmitting any part of masseter; angle of mandible remarkably
expanded transversely ; seaphoid and lunar not separate; fibula not
articulating with caleaneum, free from tibia; body stout, about 350
millimeters long; pelage dark; eyes and ears small (pl. 29) ; mammae
six; tail very short; nocturnal, fossorial, found in restrieted area in
western North America.
The much modified hyposodont teeth, the depressed skull, the
absence of postorbital processes (pls. 26, 27), the extraordinarily
inflected angle of the mandible, the fusion of the scaphoid and lunar
into a single bone, the proportionate enlargement of the forefeet (for
digging), and probably also the smallness of eyes and ears, are sug-
gestive of a high degree of specialization in Aplodontia. On the other
hand the simplicity of the dental formula and the characters of the
infraorbital region (pl. 27) show a generalized or primitive condi-
tion. The infraorbital foramen is moderate in size, and does not
transmit any portion of the masseter, the origin of which, according
to Matthew (1910, p. 69), is wholly behind and below the lower
margin of the orbit, and not extended forward on the side of the
muzzle.
3. VARIATIONS IN THE GENERIC NAME
Palmer (1904, pp. 24, 25) has ealled attention to the fact that the
name Aplodontia is capable of twenty-four modifications, each one
progressively differing from the next by a single letter. That several
of these possible spellings of the name have found place in literature
is indicated by the following list of names applied to the genus:
Anisonyx (part) Rafinesque (1817), p. 45.
Anisonix (part) Minding (1829), p. 86.
Aplodontia Richardson (1829a), pp. 333-337.
Apludontia Fischer (1830), p. 398 (error for 598).
Haplodon Wagler (1830), pp. 4, 22.
1918] Taylor: Revision of the Rodent Genus Aplodontia 453
Arctomys (part), Douglas (1914), pp. 59, 156.
Apluodontia Richardson (1837), p. 150.
Aplodontie Gervais (1854), p. 364.
Haploodon and Hapludon Brandt (1855), p. 150, footnote.
Haploodon, Haploudon, Haploddus, Haplodus, Haploudus Coues (1877b),
p- 556.
Haplodonta Cope (1883), p. 55.
Hapludus, Aploudontia, Haploudontia Coues (1890), p. 2712.
Haplodontia Elliot (1899), pp. 241-276.
Aploodontia, Aplodon, Aploodon, Aploudon, Apludon, Aplodus, Aploodus,
Aploudus, Apludus, Haploodontia, Hapludontia Palmer (1904), p. 25.
There has been no agreement as to a vernacular name for aplo-
dontia. Among names applied to the animal are found the following:
Mountain beaver, mountain boomer, gehalis, and farmer; giant mole,
mammoth mole, ground bear, marmot, ground hog, woodchuck,
gopher, badger, muskrat, blue muskrat, high ground muskrat, and
(Indian names) sewelel, shewelel, sewewel, or sewellel, showhurll,
showhurtl, showtl, showt’l, show’tl, shote, squallah, swak-la, o-gwal-lal,
and ou-ka-la. I am indebted to Dr. C. Hart Merriam for the following
list of names given aplodontia by certain tribes of Californian Indians:
NAME TRIBE LocaLiry
Ne-ta-te Tolowa or Huss Crescent City
Mah-pe-neetch Karok Happy Camp to Weitzpek
Tabt-ka-wer-itl Soo-lah-te-luk Humboldt Bay
Waw-kaw-see Mo-des-se Pit River
Yah-sah Nis-se-nan Colfax and Placerville to Yuba
Wes-skap* Yurok Lower Klamath; mouth of river
to Weitzpek
4. List of SPECIES AND SUBSPECIES WitH TyPE LOCALITIES
’
Aplodontia rufa rufa (Rafinesque). ‘‘ Neighborhood of the Columbia River”’;
specimens from Marmot, Clackamas County, Oregon, regarded as typical.
(p. 454)
Aplodontia rufa olympica Merriam, C. H. Quiniault Lake, Chehalis County,
Washington.
(p. 460)
Aplodontia rufa columbiana Taylor. Roab’s Ranch, near Hope, British
Columbia.
(p. 463)
Aplodontia rufa rainieri Merriam, C. H. Paradise Creek, south side of
Mount Rainier, Washington.
(p. 465)
Aplodontia rufa pacifica Merriam, C. H. Newport, mouth of Yaquina Bay,
Lincoln County, Oregon.
(p. 467)
Aplodontia rufa humboldtiana Taylor. Carlotta, Humboldt County, Cali-
fornia.
(p. 470)
454 University of California Publications in Zoology — (Vou. 17
Aplodontia rufa californica Peters. The mountains of California; assumed
to be the Sierra Nevada; specimens from Blue Canyon regarded as typical.
, (p. 473)
Aplodontia rufa nigra Taylor. Point Arena, Mendocino County, California.
(p. 479)
Aplodontia rufa phaea Merriam, C. H. Point Reyes, Marin County, Cali-
fornia.
(p. 480)
5. ACCOUNTS OF SPECIES
Aplodontia rufa rufa (Rafinesque)
Brown Aplodontia
Sewellel Lewis and Clark (1814), p. 176.
Sewewell [= Sewellel] Lesson (1827), p. 240.
Ground Rat Douglas (1836), p. 101.
Arctomys brachyurus? Douglas (1836), p. 101.
Anisonya? rufa Rafinesque (1817), p. 45.
Anisonyx? rousse Desmarest (1822), p. 330.
“*Anisonia rufa?’’ Minding (1829), p. 86.
Arctomys rufa, Harlan (1825), pp. 308-309.
Anisonyx roux Lesson (1827), p. 240.
Aplodontia leporina Richardson (1829a), pp. 888-337.
Apludontia leporina, Fischer (1830), p. 398 [= error for 598].
H{aplodon]. leporinus Wagler (1830), p. 22.
Apluodontia leporina Richardson (1837), p. 150.
Aplodontie léporine Gervais (1854), p. 364.
H{aploodon]. leporinus, Lilljeborg (1866), p. 41.
Haplodon rufus Coues (1877b), p. 557.
Haplodonta rufa, Cope (1883), p. 55.
Aplodontia rufa, Merriam (1886), pp. 312-328.
A[plodontia]. rufus, Price (1894), p. 328.
Aplodontia major (part) Merriam (1897), p. 219.
H{aplodontia]. rufa, Elliot (1899), p. 251.
Aplodontia chryseola Kellogg (1914), pp. 295-296.
Aplodontia rufa grisea Taylor (1916c), pp. 497-499.
Type Locality—It is well known that Rafinesque’s deseription of
“*Anisonyx? rufa’? was based entirely upon the Sewellel of Lewis
and Clark. As cited by Rafinesque the type locality is the ‘‘neigh-
borhood of the Columbia River.’’ It is now known that two subspecies
of Aplodontia occur on the Columbia River. The coastal form
deseribed as Aplodontia pacifica is found at Astoria, while an inland
form commonly known as A. rufa oceurs in the foothill country about
Mount Hood. The original description of rwfa is not diagnostic as
between the coastal and the inland forms. Doubtless Lewis and Clark
saw Indian robes made from the skins of both. There would be no
mes.
1918 ] Taylor: Revision of the Rodent Genus Aplodontia 455
advantage in transferring the name rufa to the well known form
pacifica. It seems altogether appropriate, on the other hand, that
the name rufa be fixed on the inland race, of which specimens col-
lected at Marmot, Clackamas County, Oregon (western slope of Mount
Hood, not far from the Columbia River), may be regarded as typical.
Specimens Examined—Total number, 135, from the following
localities :
British Columbia: New West-
minster Provineial Distriet—Chilli-
wack, 5; Sumas, 1; Mt. Baker
Range, 1.
Washington: Skagit County —
Sauk, 8 (1 skull only); Hamilton,
1; Mt. Vernon, 5 (2 skulls only).
King County—Seattle, 6 (one is
skull and skeleton only); Ravenna,
near Seattle, 1; Renton, near Se-
attle, 3; Kirkland, 6; ‘‘ Puget
Sound’’?, 2. Kittitas County —
Easton, 8. Pierce County—Puyal-
lup, 2.
Oregon: Clackamas County —
Marmot, 6 (2 skulls only); Eagle
Creek, eight miles southeast of
Bissell, 1; Bissell, 1; head of Hagle
Creek, 1; ‘‘Clackamas County,’’ 1
(no skull). Lane County—Vida, 6;
McKenzie Bridge, 3; Horse Pasture
Mt., ten miles by road southeast
of McKenzie Bridge, 3; O’Leary
Mt., ten miles by road southeast of
McKenzie Bridge, 1. Jackson
County—Siskiyou, 20; north base
Ashland Peak (alt. 5,200 ft.), 2.
Klamath County — Mt. Mazama,
Anna Creek (alt. 6,000 ft.), 3; Fort
Klamath, Anna Creek Canyon, 3.
California: Siskiyou County —
Siskiyou Mts., White Mt. (alt.
6,000 ft.), 8; Siskiyou Mts., Craigy
Peak (alt. 6,200 ft.), 1; Siskiyou
Mts., Studhorse Canyon (alt. 6,500
ft.), 1; Trinity Mts., east of Hoopa,
ten miles west of Forks of Salmon
(alt. 5700-5,800 ft.), 5; Salmon
Mts., Etna Mills, 3; Salmon Mts.,
Jackson Lake, 5; Salmon Mts., Wild
Cat Peak, 1; Salmon Mts., South
Fork of Salmon River, 1. Trinity
County—Canyon Creek, 5; Salmon
Fig. H. Geographic distribution of
the nine recognized races of the genus
Aplodontia.
. Aplodontia
. Aplodontia
. Aplodontia
. Aplodontia
Aplodontia
. Aplodontia
. Aplodontia
. Aplodontia
. Aplodontia
SOND wre
rufa
rufa
rufa
rufa
rufa
rufa
rufa
rufa
rufa
rufa
columbiana
olympica
rainieri
pacifica
humboldtiana
californica
nigra
phaea
456 University of California Publications in Zoology — (Vou. 17
Mts., head of Grizzly Creek, 3. Humboldt County—‘‘ Northwest California’’
| Hoopa Valley], 2 (no skulls); Rio Dell, 1.
Geographic Range.—Neighborhood of the Columbia River, in west-
ern Oregon, interiorly on the Pacific side of the Cascades; thence
southward in a belt of unknown width to Mount Mazama in southern
Oregon and the Siskiyou-Trinity district im northern California ;
northward to Puget Sound and the Chilliwack-Sumas region in south-
western British Columbia. Altitudinal range, from sea level in the
Puget Sound district to 6,500 feet in the Siskiyou-Trinity Mountains
of northern California; zonal range, Transition and Canadian.
Cranial Characters —Skulls moderate (see measurements below),
nasals variable in outline, broad anteriorly, but usually much nar-
rowed at their posterior ends; rostrum short; audital tubes in typical
material tending to be of greater caliber than in other forms of the
genus, about as in californica (see Remarks, below).
External Characters—Above, in specimens taken at all seasons,
light ochraceous-buff (Ridgway, Color Standards and Color Nomen-
clature, 1912), in some specimens grading anteriorly into light buff,
vinaceous-cinnamon or even occasionally approaching tawny; some
with a distinetly grayish, others a distinctly brownish cast of color-
ation; some examples paler anteriorly and grayer posteriorly, with
deepest coloration in middle region of body; grizzled with more or
fewer blackish hairs; white spot at base of ear; some silvery-white
hairs insprinkled, particularly posteriorly. Below, grayish, with the
faintest possible wash of brownish of a hue near pinkish buff, or
brownish, with a conspicuous wash of vinaceous-cinnamon or light
vinaceous-cinnamon, and with seattered black hairs insprinkled ; prae-
tically all hues between these are observable; drab-gray basal color-
ation showing through to a greater or less degree; spot-marks about
mammae in spring and fall females seal brown or paler in coloration.
Molt and Seasonal Change—That Aplodontia rufa rufa molts
once, that the pelage renewal takes place during the late summer and
the fall, and that a term of from two to three months is required for
the completion of the molt, is suggested by the following facts:
Specimens taken May 17, June 5, 22, 23, and July 22 and 23, and
even two exceptional specimens taken August 26 and September 13,
are much worn but show no sign of molt. On the other hand, molt
has already begun in other specimens of rufa taken on June 23 and
24, and pelage renewal is going on in practically all specimens taken
during August, September and October. Examples secured Septem-
1918 | Taylor: Revision of the Rodent Genus Aplodontia 457
ber 29 and October 9, respectively, show the last stages of the process;
and the molt is entirely completed in a specimen taken November 21.
Evidently August, September and October are the principal months
of molt.
Seasonal changes are trifling. The fresh pelage is longer and
tends to be somewhat browner, sometimes blacker, than in its hold-
over stage in spring and early summer.
CRANIAL MEASUREMENTS OF Aplodontia rufa rufa (13 SKULLS) From Marnor,
VIDA AND VICINITY OF MCKENZIE BRIDGE, OREGON
Average Mean Maximum Minimum
Basilar length 61.1 57.2
Length of nasals (10 skulls) —. 27.3 23.0
Width of nasals 13.3 11.5
Length of audital tube (10 skulls) . 21.0 16.0
Length of incisive foramen 8.7 5.7
Zygomatie width 59.6 49.5
Mastoid width of cranium ............ 59.0 47.4
Alveolar length superior cheek teeth 119°9 18.2
Distance between infraorbital formamina ~.... 16.4 17.8 15.0
Mandible, transversely across angular process... 23.1 22.4 25.0 ie)
Mandible, greatest length (12 skulls) .......... 46.2 46.0 48.7 43.5
EXTERNAL MEASUREMENTS OF Aplodontia rufa rufa, FROM VIDA AND
I
VICINITY OF MCKENZIE BRIDGE, OREGON
Average Mean Maximum Minimum
Total length (9 skims) ~............ 345 348 387 310
letihayel SHoyoyr. (C7 fbr sis), eeepc eee eae 56 56 59 54
CRANIAL MEASUREMENTS OF Aplodontia rufa rufa (12 SKULLS), FROM THE VICINITY
oF SEATTLE AND Mount BAKER RANGE, WASHINGTON, AND THE
CHILLIWACK-SuMAS District, BRITISH COLUMBIA
Average Mean Maximum Minimum
Basilar length 59.6 59.5 62.1 57.0
Length of nasals (10 skulls) 25.2 27.2 23.2
Width of nasals 11.5 12.8 10.2
Length of incisive foramen ell 8.1 6.2
Zygomatic width ............ Peeters 55.5 54.1 58.9 49.3
Greatest width of interpterygoid fossa .......... 5.0 5.0 5.8 4.2
Mastoid width of cranium ~................... 25222 52.5 55.7 49.4
Alveolar length of superior cheek teeth geil 20.0 18.3
Distance between infraorbital foramina 16.4 17.6 15.2
Mandible, transversely across angular process... 23.0 25.9 20.2
Mandible, greatest length 47.6 47.3 49.5 45.1
EXTERNAL MEASUREMENTS OF Aplodontia rufa rufa (9 SKINS), FRoM THE VICINITY
OF SEATTLE, WASHINGTON, AND CHILLIWACK, BRITISH COLUMBIA
Average Mean Maximum Minimum
Total length ... 332 345 318
Hind foot 56 58 54
458 University of California Publications in Zoology (Vou. 17
CRANIAL MEASUREMENTS OF Aplodontia rufa rufa (7 SKULLS), FRoM TRINITY
AND SISKIYOU COUNTIES, CALIFORNIA
Average Mean Maximum Minimum
Basilar length 59.3 59.8 61.8 57.9
Length of nasals (6 skulls) .. 24.5 26.2 22.9
Width of masals) ---------- 11.8 13.0 10.7
Length of audital tube .... 19.1 19.4 21.1 17.7
Length of incisive foramen ws Ue 7.2 7.6 6.7
Zi; FOU ULC avy Cit Np ae sees es neces tae sees teat rece oee stoner 56.1 55.9 60.4 51.4
Greatest width of interpterygoid fossa —....... 5.1 5.1 5.3 4.8
Mastoid width of cranium ....................... 3.6 54.3 58.2 50.4
Alveolar length superior cheek teeth --- 18.8 18.7 19.2 18.2
Distance between infraorbital foramina ...... 16.5 16.3 18.2 14.4
Mandible, transversely across angular process... 24.0 23.9 26.0 21.7
Mandible, greatest length —......-.-.---------------: 48.3 enor 51.3 44.5
EXTERNAL MEASUREMENTS OF Aplodontia rufa rufa (7 SKULLS), From TRINITY
AND SISKIyoU COUNTIES, CALIFORNIA
Average Mean Maximum Minimum
340 370 310
59 58 63 53
Total length
Elinid! footie.
Averages of External Measurements: Four specimens from Easton, Wash-
ington: total length, 354 mm. (345-368); hind foot, 59 (58-63). Eighteen
specimens from Siskiyou, Oregon: total length, 332 (310-394); hind foot, 58
(52-63). Ten additional examples from the Siskiyou Mountains, California:
total length, 349 (320-370); hind foot, 55 (53-58). Two specimens from Ash-
land Peak, Oregon: total length, 340, 352; hind foot, 60, 63. Two examples from
Anna Creek, Mount Mazama: total length, 360, 385; hind foot, 58, 61. Three
specimens from Fort Klamath, Anna Creek Canyon: total length, 333, 338, 370;
hind foot, 54, 60, 57. Three specimens from Canyon Creek, California: total
length, 345, 360, 370; hind foot, 60, 60, 60.
Remarks —Aplodontia rufa rufa is a variable form embracing a
number of local forms, some of which, if their extremes only were
considered, would certainly rank as good subspecies. The number
of these local variants (which have been noted from the Chilliwack-
Sumas district, British Columbia; vicinity of Seattle, Puget Sound,
Washington; Ashland Peak, Siskiyou and Fort Klamath, Oregon; the
Siskiyou Mountains and Canyon Creek, California; and elsewhere),
coupled with the great range of individual variation observable in
series from the same general locality, make impracticable the recogni-
tion of subspecies within the range of A. r. rufa as here outlined.
Specimens of rufa from the immediate vicinity of Seattle, Wash-
ington, tend to have narrower nasals and rostra, mastoid width less,
audital tubes of smaller caliber, and fossae set off anteriorly by lamb-
doidal ridge shallower than in typical material from Marmot, Oregon.
1918] Taylor: Revision of the Rodent Genus Aplodontia 459
Externally they give a mass effect of grayish rather than brownish
as in the Marmot examples. The differences are slight, however, and
are not shown to the same extent by examples from nearby localities,
as Mount Vernon, Hamilton, Sauk and Easton, in Washington, and
the Chilliwack-Sumas district in British Columbia. The form from
the vicinity of Seattle was recently described by the writer under the
name Aplodontia rufa grisea (Taylor, 1916c, p. 497), but the exam-
ination of considerable additional material indicates the propriety of
synonymizing grisea under rufa.
The effects of varying degrees of local isolation seem to be shown
by specimens of Aplodontia rufa from the rough and mountainous
region embraced in its range in southern Oregon and northern Cali-
fornia. Groups of specimens from particular localities, however, do
not show the constant differences which would be necessary to entitle
them to recognition as subspecies. Examples of rufa from the Siski-
you-Trinity region of northern California (recently deseribed by
Kellogg, 1914, p. 295, as Aplodontia chryseola) tend to have the eali-
ber of the audital tube less than in typical rufa, the outline of the
external auditory meatus a little flatter, the nasals slightly narrower
and shorter, and coloration and quality of pelage slightly different.
These tendencies, however, are overshadowed by the magnitude of the
individual variation in the series. There appear to be several other
local races of rwfa as well entitled to subspecifie recognition as chry-
seola, but if they were to be described, no logical ranges could be given,
and the degree of overlapping would make identification of specimens
difficult if not impossible.
Intergradation between Aplodontia rufa rufa and neighboring
subspecies is hinted at or directly demonstrated by specimens
examined as follows: Intergradation with columbiana, by specimens
from Sumas and Chilliwack, British Columbia; with rainieri, by
examples from Easton, Washington; with olympica, by an example
from Steilacoom, Washington; with pacifica, by specimens from Lane
County, Oregon, and from Siskiyou, Oregon; with californica, by
examples from Mount Mazama, Oregon, and Canyon Creek, Cali-
fornia; and with humboldtiana by specimens from the divide between
the Trinity and Klamath rivers, twelve miles north of Hoopa Post
Office, northern California.
A considerable series of specimens from Siskiyou, Oregon, is puz-
zling, being pacifica-like in coloration, but larger, and in certain
cranial characters more like rufa. As in other series, the range of
460 University of California Publications in Zoology — (Vou. 17
individual variation is great. Two examples from Siskiyou (nos. 56737,
56738, Biol. Surv. Coll.) are larger than the others of the series and
have conspicuously heavier crania. Examples of rufa from Ashland
Peak, also, are above the average in size, and show a likeness to rainieri
in certain skull characters.
Intergradation between rufa and californica is best shown by sev-
eral examples from Canyon Creek, California. These specimens com-
bine the coloration and shortness of the rostrum of rufa with the
squarer zygomatic arches and, in one ease, the distinctive nasal out-
line of californica.
The distribution of aplodontias in northwestern California will
bear additional intensive research. Specimens from the Trinity
Mountains, east of Hoopa, are nearly typical of rufa from farther
inland, although they show a slight tendency eranially in the direction
of humboldtiana. Real intergradation is indicated, however, by three
specimens from the divide between the Klamath and Trinity rivers,
twelve miles north of Hoopa Post Office. Of these, an adult female
(no. 98745, Biol. Surv. Coll.) both in eranial and external characters
resembles humboldtiana; one young animal also (no. 97291, Biol. Surv.
Coll.) resembles humboldtiana; while the third (no. 97290, Biol. Surv.
Coll.), also a young animal, is nearest rufa. Examples referred to rufa
have been taken in Hoopa Valley, as well as at Rio Dell, Humboldt
County, California.
A tendency toward pattern formation is noted in specimens from
Sauk, Washington. Extensive irregular patches of white beneath are
conspicuous in several examples.
Aplodontia rufa olympica Merriam
Olympie Aplodontia
Aplodontia olympica Merriam (1899), p. 20.
Haplodontia olympica, Elliot (1899), pp. 251-253.
Type.—Male, young adult, no. 89549, U. S. Nat. Mus., Biol. Surv.
Coll.; Quiniault Lake, Olympic Mountains, Washington; July 24,
1897 ; collected by R. T. Young; orig. no. 309; stuffed skin, with skull
and jaws, all in good condition.
Specimens Examined.—Total number 29, from the following
localities :
1918 | Taylor: Revision of the Rodent Genus Aplodontia 461
Washington: Pierce County—Fort Steilacoom, 1; Steilacoom, 1 (skull only).
Clallam County—Happy Lake, Olympie Mts., 6; Olympie Mts., near head of
Soledue River (alt. 4500 ft.), 1; Port Angeles, 3. Jefferson County—Olympie
Mts., head of north fork of Skokomish River, 2. Chehalis County—Quiniault
Lake, 8. Mason County—Lake Cushman, 5; ‘‘Mason County,’’ 2 (odd skulls).
Geographic Range—Northwestern Washington, vicinity of Olym-
pic Mountains, intergrading with Aplodontia rufa rufa in the vicinity
of Steilacoom, southern Puget Sound.
Cranial Characters——Skull moderate in size (see measurements
below) ; nasals tending to be somewhat variable in outline; temporal
lnes or ridges tending to be more closely approximated anteriorly
than in rufa; zygomatic arches tending to be lighter in weight than
in rufa; thirteen out of seventeen crania examined with prominent
postorbital process on the jugals, a character not developed to the
same degree in any other form of Aplodontia; audital tubes tending
to be of smaller caliber than in typical rufa; notch on upper side of
external auditory meatus smaller and narrower than in typical rufa.
External Characters—Above, 1 summer specimens, pinkish cinna-
mon to light ochraceous-buff, often with something of a grayish cast ;
the whole grizzled with more or less of an admixture of black hairs;
head and face brownish or grayish; under parts grayish with a faint
wash of pinkish buff or cinnamon buff.
Sexual Differences—No appreciable external differences due to
sex have been observed except in respect to the spot markings of the
mammae and possibly a slight pallor ventrally in females.
CRANIAL MEASUREMENTS OF Aplodontia rufa olympica (17 SKULLS)
Average Mean Maximum Minimum
Basilar length 2 09:3 60.2 63.0 57.4
Length of nasals (12 skulls) . 26.4 26.9 28.9 24.9
Wadithwotemiasallsi-e--eeeee TES 12.1 13.3 10.8
Length of audital tube (12 skulls) - 18.9 19.1 20.0 18.1
Length of incisive foramen. .... 7.0 7.2 8.0 6.4
Zygomatic width (15 skulls) 54.6 54.9 56.9 52.8
Greatest width of interpterygoid fossa (12 skulls) 5.2 5.2 6.0 4.4
Mastoid width of cranium (16 skulls) —- ee 05.0) 52.8 55.9 49.7
Alveolar length of superior cheek teeth 19.3 19.4 20.2 18.5
Distance between infraorbital foramina Bee Gto 16.4 17.5 15.3
Mandible, transversely across angular process ... 23.3 23.5 25.2 21.9
Mandible, greatest length ...............--------------------+ 47.1 46.8 49.7 43.8
EXTERNAL MEASUREMENTS OF Aplodontia rufa olympica (10 SKINS)
Average Mean Maximum Minimum
355 353 367 340
49 47 59 36
Total length
Hind foot
462 Umversity of Califorma Publications in Zoology — (Vou. 17
Molt and Seasonal Change—The only months represented by
available specimens of olympica are July and August. In the series
at hand the molt begins about the middle of July, and is not com-
pleted till September at the earliest. One example (no. 67615, 9,
U.S. Nat. Mus., Biol. Surv. Coll.), secured August 17, is exceptional
in that even at that late date it shows no sign of molt. On the basis
of specimens at hand the winter coat is a little browner than the
summer pelage.
Figs. I-K. Side view of crania of Aplodontia rufa olympica, to illustrate
variation in development of postorbital process of the jugal. XxX 1. Fig. I, no.
66229, Biol. Sury. Coll.; fig. J, no. 6314, Field Mus. Nat. Hist.; fig. K, no. 89549,
Biol. Sury. Coll.
Remarks.—Aplodontia rufa olympica is a feebly marked race of
the rufa series occurring in the Olympic Mountain district of Wash-
ington, and grading insensibly into rufa to the east and southeast.
Additional material from scattered localities in western and north-
western Washington may demonstrate so great a degree of variability
as to make inadvisable the separate recognition of the Olympic Moun-
tain form. The postorbital process on the jugal, the most important
1918 | Taylor: Revision of the Rodent Genus Aplodontia 463
character of the subspecies, is subject to considerable variation (text
figs. I, J, K), and is more apparent in specimens from Lake Cushman
than in those from Quiniault Lake. Specimens from Steilacoom (no.
2476, U.S. Nat. Mus., skull only) and Fort Steilacoom (no. 278, U. 8.
Nat. Mus., part of skull inside skin), here referred to olympica, might
be referred to rufa with equal propriety.
Aplodontia rufa columbiana Taylor
Northern Aplodontia
Aplodontia californica columbiana Taylor (1916c), pp. 499-501.
Type.—Male adult ; no. 1899, Coll. E. A. and O. Bangs, Mus. Comp.
Zool.; Roab’s Ranch, Hope, British Columbia; June 14, 1894; col-
lected by W. C. Colt; stuffed skin with skull and jaws in good econdi-
tion, except skin of foreleg injured in trap, and skull with left audital
tube, region of foramen magnum, and hamulars, somewhat injured.
Geographic Range.—Vicinity of Hope, British Columbia, south in
the Cascade Mountains of Washington; probably intergrading with
Aplodontia rufa rainiert between the international boundary and
Mount Rainier.
Specimens Examined—Total number 11, from the following
localities :
British Columbia: Yale Provincial District—Lake House, near Hope, 4;
Roab’s Ranch, Hope, 5. Washington: Skagit County—Head of Cascade River, 2.
Cranial Characters—Skull heavy and large (see measurements
below) ; for example, on basis of tables of measurements, furnishing
maximum for the genus in length of nasals, zygomatic width, mastoid
width, and greatest length of mandible; averages of length of nasals
and zygomatic width exceeding the maximum in any other form; zygo-
matic arches moderate and, looking down upon them from above, trans-
versely expanded posteriorly ; temporal lines or ridges tending to be
closely approaching, though never forming a distinct sagittal crest ;
dorsal outline of skull comparatively straight; audital tubes small
ealibered, dorso-ventral diameter tending to exceed anteroposterior
diameter, tube thus appearing as if pinched anteroposteriorly.
External Characters—Above, in summer skins, near light pinkish
cinnamon, in some specimens as pale as light ochraceous-buff, in others
as dark as pinkish cinnamon, grizzled with many blackish and some-
times a few whitish-tipped hairs; number and degree of concentration
of black hairs on back variable ; underparts showing faint brown wash,
464 University of California Publications in Zoology — [Vow. 17
sometimes nearer light pinkish cinnamon, sometimes nearer pinkish
buff; general undertone of coloration ventrally pale drab-gray, with
an insprinkling of varying numbers of blackish, whitish, cinnamon or
buffy hairs; irregular areas of hair white from base to tip appearing
in most of the males ventrally.
Sexual Differences.—The conspicuous white patches ventrally ap-
pear in males only, all the examples of that sex showing them to a
greater or less extent. The spot marks about the mammae in May and
July females are less conspicuous than usual. Of two females taken
in July, one (no. 1894, Mus. Comp. Zool.) has the hair about the
mammae practically worn off, while the other (no. 1895, Mus. Comp.
Zool.) has the hair in the same region unworn. It seems likely that
the condition of the hair about the mammae may be taken as an index
to breeding activity.
CRANIAL MEASUREMENTS OF Aplodontia rufa columbiana (9 SKULLS)
Average Mean Maximum Minimum
Wen oth sorte nasallse (/ees ks) pcos eee eee 29.3 29.5 30.9 28.2
Width of nasals 12.8 12.9 13.8 12.0
Length of incisive foramen 8.0 7.8 8.5 1.2
Zygomatic width 62.3 60.9 64.0 57.9
Mastoid width ‘of eramiumy 2cete ete ceereee cree 57.5 57.2 61.2 53.2
Alveolar length superior cheek teeth . 19.3 19.3 19.8 18.8
Distance between infraorbital foramina 16.9 17.2 18.3 16.2
Mandible, transversely across angular process .... 24.6 24.5 26.1 23.0
Mandible ereatest Lemp tl) 22sec cre eee 51.2 51.9 54.9 48.9
EXTERNAL MEASUREMENTS OF Aplodontia rufa columbiana (9 SKINS)
Average Mean Maximum Minimum
Mortal len orthig ecco 427 425 470 380
Remarks.—Available material indicates intergradation between
Aplodontia rufa columbiana and A. r. rufa on the one hand and A. r.
rainiert on the other. Specimens in the United States National Museum
. : 30739 3089
from the head of the Cascade River, Washington (nos. 22732 30890
426399
42790’
Biol. Sury. Coll.) are intermediate between columbiana and rainieri,
although perhaps a trifle closer to the former. If, as seems likely, the
distribution of Aplodontia is more or less continuous between Mount
Rainier and the mountains in the Hope District, British Columbia, it
is probable that one grades into the other by insensible degrees. Aside
from its greater dimensions columbiana may be separated from rufa
through its smaller-calibered, more pinched-up audital tubes, as com-
pared with the larger, rounder tubes of rufa. Columbiana also has the
zygomatie arch more expanded posteriorly than in rufa, as well as
broader nasals posteriorly, and more closely approximated temporal
1918] Taylor: Revision of the Rodent Genus Aplodontia 465
ridges. From rainicrt typical columbiana differs in having eranial
measurements averaging greater, the difference being shown in length
and width of nasals, length of incisive foramina, zygomatie width,
mastoid width, and greatest length of mandible. The external auditory
meatus tends to be of different shape in columbiana, being pinched up
anteroposteriorly, while in rainier?i the meatus is rounder and tends to
be flattened dorsoventrally.
Aplodontia rufa rainieri Merriam
Mount Rainier Aplodontia
Aplodontia major rainieri Merriam (1898a), p. 21.
Haplodontia rufa rainieri, Elliot (1901), p. 112.
A[plodontia]. r[ufa]. rainieri, Trouessart (1904), p. 348.
Type—Male adult, skull and skin; no. 90144, U. S. Nat. Mus.,
Biol. Sury. Coll.; Paradise Creek, south side of Mt. Rainier, Wash-
ington, alt. 5200 ft.; Aug. 6, 1897; collected by Vernon Bailey; orig.
no. 6122.
Specimens Examined.—Total number 9, all from Washington ;
Pieree County—Mt. Rainier, Paradise Creek, 9 (1 skull only).
Geographic Range.—Known only from yieinity of type locality.
Cranial Characters —Skull large (see measurements below) ; nasals
long and comparatively straight sided; posterior two-thirds of outline
tending to be shghtly convex laterally, but usually with a shght lateral
embayment far back; rostrum broad, as in columbiana; zygomatic
arches moderate, proportionally lighter in weight, not so much ex-
panded near posterior root as in columbiana, flat beneath in vicinity
of posterior root; temporal ridges tending to remain separated for
their entire length by several millimeters; caliber of audital tubes
small; external auditory meatus round or flattened dorsoventrally.
External Characters—Above, in summer skins, light ochraceous-
buff, grizzled with blackish, the black hairs often with silvery white
tips; a tendency observable toward concentration of blackish on middle
line of back; sides having comparatively few black hairs; underparts
deep quaker drab to light quaker drab, marked with whitish generally,
such markings more conspicuous anteriorly on the throat; spot marks
near light brownish drab; a sparse insprinkling of blackish hairs; a
faint wash of pinkish buff in some examples.
Age Variation —There is not so striking an actual decrease in
width of interorbital constriction with age as is usual in the genus.
466 University of California Publications in Zoology — [Vou. 17
The youngest specimens all have this measurement broad. In some
of the older ones it is broad, in others narrow. The zygomata are
usually squarer and more expanded in the adults than in the young.
The mastoid process is exceeded by the audital tube in all the adults,
being, however, only slightly exceeded in the oldest ones, while in
young specimens the audital tube materially exceeds the mastoid
process. The width of the palate between the third premolar and the
incisive foramina remains about the same, or even increases slightly,
with age. Looking at the crania in side view, the ventral outline of
the rostrum is nearly plane in the young, with slight eminences ob-
servable at the posterior end of the incisive foramina. In the adults
the outline tends to be more rounded, the slight eminences having
disappeared.
Molt.—Specimens collected on Mount Rainier on August 6 and 7
are with one exception (no. 90143, 9, U.S. Nat. Mus., Biol. Surv. Coll.)
beginning to molt. In one example (no. 90145, 9, U. S. Nat. Mus.,
Biol. Surv. Coll.) hair renewal is taking place on small areas on the
sides; in another (no. 90144, 4) new hair is coming in on the right
side only on an area one inch wide and four inches long, as well as
in an irregular area on the top of the head; still another (no. 90137, ¢)
is molting extensively laterally as well as in a small area on the head.
The condition of this specimen suggests that the molt begins on the
sides, and sometimes becomes quite extensive before the head molt
starts at all.
CRANIAL MEASUREMENTS OF TYPE AND TOPOTYPES OF Aplodontia rufa rainieri
(4 SKULLS)
Average Mean Maximum Minimum
Basilar length 62.1 63.6 65.4 61.9
Length of nasals (3 skulls)
Width of nasals 12.7 14.0 11.4
Length of audital tube (2 skulls) 19.1 19°9 18.4
Length of incisive foramen .-........... tlstf 8.2 7.2
iy PROAEKOSAMG UTE | sone toe en smack anes one eace aos : 58.6 62.0 55.3
Greatest width of interpterygoid fossa 5.6 5.9 5.4
Mastoid width of cranium 55.6 57.8 53.5
Alveolar length superior cheek teeth —. 19.9 21.0 18.9
Distance between infraorbital foramina : . Ufeil 18.1 16.2
Mandible, transversely across angular process ... 25.7 26.0 28.2 23.9
Mandible, greatest length —...--.------2.---.2---te-=---- 50.8 50.8 52.9 48.8
EXTERNAL MEASUREMENTS OF TyPE AND ToporTyPES oF Aplodontia rufa rainieri
(4 Skins)
Average Mean Maximum Minimum
Total length 377 379 384 365
TBnC GGL AKO YO poets cee ce 62 63 65 62
1918 ] Taylor: Revision of the Rodent Genus Aplodontia 467
Remarks.—Comparisons of examples of Aplodontia rufa rainicri
from the type locality and A. r. columbiana from the head of the
Cascade River, Washington, with specimens of A. r. rufa from the
Puget Sound district (Kirkland, Seattle, Chilliwack, Sumas), and
then with examples from localities more or less intermediate geograph-
ically and altitudinally, as Sauk and Easton, Washington, demonstrate
beyond a doubt that intergradation takes place between A. r. rufa
and the rainieri-columbiana series. A gradual change is indicated
from the lowland rufa type to the mountain rainieri-columbiana type
in size, in general cranial characters, and, particularly, in the dimen-
sions and outline of the nasal bones and the size and outline of the
external auditory meatus. More abundant material would doubtless
supply more complete evidence for intergradation.
A. r. rainieri tends to be grayer than A. r. rufa, decidedly grayer
than in the typical form, and to have less brown beneath. The nasals
in rainiert tend to be broader posteriorly, and the caliber of the audital
tubes tends to be less.
Aplodontia rufa pacifica Merriam
Pacific Aplodontia
Aplodontia pacifica Merriam (1899a), p. 19.
Haplodontia pacifica, Elliot (1901), p. 114.
Type—Female adult, skull and skin; no. 77372, U. S. Nat. Mus.,
Biol. Surv. Coll.; Newport, mouth of Yaquina Bay, Oregon; March
20, 1896; collected by B. J. Bretherton; orig. no. 2219.
Specimens Examined.—A total of 46, from the following localities :
Oregon: Clatsop County—Astoria, 2; Mishawaka, 1 (skin only). Tillamook
County—Wilson River, MeNamer’s Camp, 1. Lincoln County—Newport, 8.
Lane County—Florence, 7; Mercer, 3; Eugene, 1; Spencer Butte, 4; Seaton, 3;
Mapleton, 3. Douglas County—Smith River, 2; Gardiner, 4. Coos County
Coquille, 3 (1 skull only); ‘‘Coos County,’’ 1. Curry County—Agness, 1; Port
Orford, 1 (skull only). Josephine County—Briggs Creek (alt. 3000 ft.), 13
miles southwest of Galice, 1.
Geographic Range.—Coast of Oregon, from Astoria on the north
at least to Port Orford on the south; ranging inland locally, as in
the vicinity of Eugene, Oregon, and gradually intergrading with
Aplodontia rufa rufa, probably in a broad belt centrally on the Pacific
slope of Oregon from the northern to the southern boundaries of the
state.
468 University of Califorma Publications in Zoology | Vou. 17
Cranial Characters ——Skull comparatively small (sée measurements
beyond) ; nasals broad anteriorly, becoming only a little narrower
posteriorly, fairly uniform in outline through the series, comparatively
straight laterally, with little or no anterior dilation; temporal lines
or ridges not strongly marked, never approximated, though sometimes
approaching to within a few millimeters of one another anteriorly ;
zygomatic arches comparatively light in weight, with weakly devel-
oped postorbital processes in certain specimens; fossae in front of
lambdoidal ridge shallow; audital tubes of small caliber, notch dor-
sally in tubes short and broad; averaging smaller, on the basis of
specimens measured, than Aplodontia rufa rufa, A. r. olympica, or
A. r. humboldtiana in basilar length, width of nasals, length of audital
tube, zygomatic width, mastoid width, alveolar length superior cheek
teeth, distance between infraorbital foramina, and greatest length of
mandible.
External Characters——Aboye, in winter specimens, sayal brown
or cinnamon to pinkish cinnamon, in one specimen (no. 9077, Field
Mus. Nat. Hist.) approximating pinkish buff; many glossy black hairs
interspersed, especially on back; top of head usually conspicuously
black ; in most specimens a black area or ill-defined broad band starting
at nose and continuing back over head and posteriorly along middle
of back to posterior end of body, this indefinite band grading into the
browner coloration of the sides; face grayish in most examples ; under-
parts grayish, with a more or less distinct brown wash varying from
cinnamon or pinkish cinnamon to warm buff or pinkish buff.
CRANIAL MEASUREMENTS OF Aplodontia rufa pacifica (23 SKULLS)
Average Mean Maximum Minimum
Basilarslentott hye eee Be ice eee 57.0 57.5 62.1 52.9
Length of nasals (19 skulls) a BASES 23.9 26.5 21.4
Wal Gh fea. caill eee ene eee eee -- 10.4 10.5 11.4 9.6
Length of audital tube (15 skulls) 16.8 16.8 17.9 15.7
Length of incisive foramen ........... cao, MTHS) 6.6 7.7 5.5
Ley feqOssTEH AKG NKR See en =) 015 51.6 56.1 47.1
Greatest width of interpterygoid fossa ............. 4.6 4.4 5.3 3.5
Mastoid width of cranium (22 skulls) —....-.... 49.1 — 50.0 52.9 47.2
Alveolar length superior cheek teeth ....... =e SiG 18.6 20.3 16.9
Distance between infraorbital foramina = 6:0 15.8 17.5 14.1
Mandible, transversely across angular process ... 22.5 23.1 25.3 20.9
Greatest length of mandible (22 skulls) ......... 45.7 45.6 49.4 41.9
EXTERNAL MEASUREMENTS OF Aplodontia rufa pacifica (23 SKINS)
Average Mean Maximum Minimum
Motall len gihye eases 316 324 356 293
Hing) footw ee 51 52 57 48
1918 | Taylor: Revision of the Rodent Genus Aplodontia 469
Molt.—The molting process is just beginning in specimens of this
subspecies taken June 19, June 29, July 17, and September 12. The
example taken on the date last named is exceptional in molting so
late. The end of the molt is shown by specimens secured on October 6
and October 24. Others taken October 5, 9, and 24 have completely
assumed the new pelage.
Remarks.—A plodontia rufa pacifica is usually separable from A. r.
olympica by both external and eranial characters. Cranially the ab-
sence in pacifica of the postorbital process on the jugal in most cases
serves to distinguish it from olympica. The nasals tend to be nar-
rower as well as shorter in pacifica, and to be straighter sided. The,
skull is usually smaller in pacifica, and the temporal lines are more
accentuated. Pacifica is grayer headed than is olympica, and there
is In pactfica a concentration of blackish on the middle line of the
back which is not so apparent in olympica. There is more brownish
dorsally in pacifica. Ventrally pacifica has a conspicuous brown wash,
while in olympica the ventral brownness is less noticeable.
From Aplodontia rufa rufa, A. r. pacifica may usually be separated
through having a greater concentration of blackish dorsally. A uni-
form brownish tone dorsally is never observed in typical pacifica, but
in rufa it is often observed. In specimens which are not so distinctly
brownish pacifica tends to be blackish while rufa tends to be grayish.
Color fails completely to allocate certain specimens from intermediate
localities. Thus examples from Siskiyou, Oregon, are in coloration
pacifica, while in size and certain skull characters they are closer to
rufa. Others from the vicinity of Eugene and other localities in Lane
County, Oregon, have the coloration of rufa but certain other char-
acters of pacifica. Cranially pacifica is usually separable from rufa
through possession of nasals narrower anteriorly and proportionally
broader posteriorly, lighter zygomatic arches, smaller average measure-
ments throughout, audital tubes shorter and of lesser caliber, and
shallower fossae anterior of the lambdoidal crests.
Smaller size and different coloration separate Aplodontia rufa
pacifica from A. r. humboldtiana. From A. r. pacifica one gets the
impression of rich brown with black hairs plentifully insprinkled and
specially emphasized on the middle line of the back, while from
humboldtiana one receives the impression of black sparsely inter-
spersed with buffy. Cranially pacifica, while very close to humboldt-
iana, can usually be separated therefrom through the possession of
nasals with straighter lateral outline.
,
470 University of California Publications in Zoology (Vou. 17
Specimens of Aplodontia rufa pacifica from Mercer, Lane County,
Oregon, tend to be uniformly larger than others from the coastal
region. One of them (no. 1600, Univ. Ore. Mus.) gives the maximum
measurement in a series of more than twenty skulls in basilar length,
length of nasals, zygomatic width, alveolar length superior cheek teeth,
distance between infraorbital foramina, distance transversely across
angular process of mandible, and greatest length of mandible.
All specimens of aplodontia collected on or very close to the coast
line (exeept the examples from Mercer just mentioned, and a skull
only, from Port Orford, Oregon, no. 206368, U. S. Nat. Mus., Biol.
Surv. Coll., which has broader nasals) are typical of the Newport
form; but as specimens from points farther away from the coast are
examined an increase in size is apparent, and the characters and color-
ation tend in the direction of rufa. To this category belong specimens
from Lane County, referred to pacifica (as no. 204887, U. S. Nat.
Mus., Biol. Surv. Coll.), examples from Siskiyou, Oregon, referred to
rufa (see p. 460), and a specimen from Briggs Creek, Oregon, thirteen
miles southwest of Galice (no. 205239, U. 8S. Nat. Mus., Biol. Surv.
Coll.), referred to pacifica. Examples from these localities partake
of the characteristics of two subspecies often in nearly the same degree,
and some or all of them might be referred to one with almost as much
propriety as to the other.
Intergradation with rufa is demonstrated by specimens of pacifica
from Spencer Butte, seven miles south of Eugene, Oregon, and by
examples of rufa from Siskiyou, Oregon. It seems probable that inter-
eradation between the two forms takes place over a broad area north
and south through central Oregon.
Aplodontia rufa humboldtiana Taylor
Humboldt Aplodontia
Aplodontia rufus (part), Price (1894), p. 328.
Haplodontia phaea (part), Elliot (1903), pp. 184, 185.
Aplodontia phaea (part), Stephens (1906), p. 95.
Aplodontia humboldtiana Taylor (1916a), pp. 21-23.
Type—Male adult; no. 21162, Mus. Vert. Zool.; Carlotta, Hum-
boldt County, California; January 4, 1914; collected by H. E. Wilder ;
orig. no. 1494; stuffed skin, with skull and jaws, all in good condition.
Specimens Examined.—A total of 24, from the following localities :
California: Del Norte County—Requa, 1. Humboldt County—Sam Lane’s
Ranch, 12 miles north of Hoopa, 3 miles southwest of Weitzpek, 3; Eureka, 5;
Carlotta, 8; Cuddeback, 7.
1918 ] Taylor: Revision of the Rodent Genus Aplodontia 471
Geographic Range—The northern coast district of California from
Humboldt Bay, Carlotta, and Cuddeback along the coast in Humboldt
and Del Norte counties northward, at least to Requa; ranging inland
locally in Humboldt County and intergrading with A. 7. rufa in the
vicinity of Weitzpek.
Figs. L, M. Dorsal view of crania of Aplodontia rufa rufa and Aplodontia
rufa humboldtiana, to illustrate different nasal outline. X 1. Fig. L, no. 13326,
Mus. Vert. Zool., Aplodontia rufa rufa, Jackson Lake, Siskiyou County, Cali-
fornia; fig. M, no. 21162, Mus. Vert. Zool., Aplodontia rufa humboldtiana, Car-
lotta, Humboldt County, California.
Cranial Characters—Skull moderate in size (see measurements
below) ; nasal outline variable, but usually broad anteriorly, with a
shallow embayment in lateral outline, not exceptionally compressed
posteriorly (see text fig. M); zygomatie arches light to medium in
472 University of California Publications in Zoology — Vou. 17
weight, with position of postorbital processes on the jugal faintly
indicated in six crania out of sixteen; temporal ridges comparatively
well marked, more accentuated than in either Aplodontia rufa rufa
or A. r. pacifica, in some skulls widely separated, as in pacifica, Mm
others approaching to within a few millimeters of each other, as in
rufa; audital tubes tending to be of smaller caliber than in rufa,
larger and straighter than in pacifica; incisive foramina comparatively
short.
External Characters.—Above, light ochraceous-buff or pinkish buff,
obscured by an admixture of black hairs which is nearly uniform over
all upper parts—an insprinkling of silvery-tipped hairs augments the
grizzled appearance—some specimens appearing conspicuously black-
ish ; under parts near pale quaker drab with many silvery white hairs;
the faintest possible wash of buffy brown observable in certain ex-
amples; a white spot usually present posteriorly in the vicinity of the
external genitalia.
Sexual Differences—Three males out of eleven have total length
greater than the maximum of that measurement in the females; six
have tail vertebrae longer than this maximum, and six also have hind
foot longer. Comparison of fall examples demonstrates that the three
darkest specimens are males, their darker coloration being more evi-
dent ventrally. In this species in fall pelage the spot marks about
the mammae do not afford a distinctive character. While they are
in evidence in certain specimens of both sexes, they are lacking in
others. In examples having the spot marks, females have them more
accentuated than males.
There is a tendency for the ridges and processes in the skulls of
males to be somewhat more accentuated than in those of females.
Temporal lines or ridges appear to come closer together with old age
in males than in females. :
Molt-—Two young specimens (nos. 9061, 9062, Field Mus. Nat.
Hist.), collected August 16 and August 18, respectively, are molting
into the adult pelage. The adult coat has replaced the juvenal pelage
everywhere in these specimens except in a narrow band across the
shoulders, although a patchy appearance posteriorly in one of them
(no. 9061) suggests that the molt is incomplete in that region. An
adult specimen (no. 21159, 4, Mus. Vert. Zool.) taken at Carlotta,
’ Humboldt County, November 5, has almost completely assumed the
fresh pelage. New hair is still coming in on a narrow longitudinal
line on the back posteriorly. Another specimen (no. 21156, ¢, Mus.
Vert. Zool.) taken October 29 has the winter pelage complete.
1918] Taylor: Revision of the Rodent Genus Aplodontia 473
CRANIAL MEASUREMENTS OF Aplodontia humboldtiana (15 SKULLS)
Average Mean Maximum Minimum
TEYYSVU Ea TTS lO ae et pee eee eee .- 58.8 58.7 55.0
Length of nasals (11 skulls) 24.5 23.8 21.9
Width of nasals —........... 11.2 11.2 10.0
Length of audital tube (12 18.4 18.0 16.5
Length of incisive foramen .- 5.9 6.5 4.8
Pi AEROVSTED ING, ASAI NE, oe eon 53.8 53.2 57.9 48.6
Greatest width of interpterygoid fossa ........... 5.5 5.7 6.7 4.8
Mastoid width of cranium .......00.0......... eee 51.7 57.4 46.0
Alveolar length superior cheek teeth ...... =. 18:8 18.6 19.7 17.6
Distance between infraorbital foramina ............ 16.4 ileal 18.9 15.4
Mandible, transversely across angular process ... 22.2 21.6 23.9 19.4
Mamaiblessoreatest Lemp th) <2... oo. 2 ccccsccsteccecescereveorns 48.1 48.1 50.7 45.6
EXTERNAL MEASUREMENTS OF Aplodontia humboldtiana (15 SKINs)
Average Mean Maximum Minimum
AN aysCell evakag a, oe ree eee reer 346 333 367 300
Ti OO teases see anecen eee 56 56 63 50
Remarks.—A plodontia rufa hwmboldtiana is larger and less richly
colored than Aplodontia rufa pacifica of the coast region of Oregon
(see p. 469). Humboldtiana is similar in coloration to A. r. rufa, of
the Trinity-Siskiyou Mountain region to the eastward, but darker, a
paler hue of the brown series of colors being interspersed with the
black hairs. The Humboldt aplodontia is not so black as is A. r. nigra,
its nearest neighbor on the south, although in certain cranial charac-
ters, notably width of nasals, length of incisive foramen, zygomatic
width, greatest width of interpterygoid fossa, and mastoid width, it
is closer to nigra than to any other one of its neighbors.
Specimens from Sam Lane’s Ranch, twelve miles north of Hoopa
and three miles southwest of Weitzpek, on the divide between the
Klamath and Trinity rivers (nos. 97290, 97291, 98475, U.S. Nat. Mus.,
Biol. Surv. Coll.), here referred to humboldtiana, show intergradation
in the direction of A. r. rufa.
Aplodontia rufa californica (Peters)
Sierra Aplodontia
Aplodontia leporina (part), Audubon and Bachman (1854), pp. 99-103.
Marmot, Mammoth Mole, Calif. Acad. Nat. Sci. (1855), p. 71.
H{aplodon]. leporinus var. Californicus Peters (1865), pp. 177-179.
H{aploodon]. californicus, Lilljeborg (1866), p. 41.
Haplodon rufus, Coues (1877b), p. 557.
Aplodontia major Merriam (1886), p. 316.
H{aplodon]. major, Townsend (1887), p. 174, footnote.
H{aplodontia]. major, Elliot (1899), p. 251.
Haplodontia rufa californica, Elliot (1901), p. 112.
Aplodontia rufa californica, Trouessart (1904), p. 348.
Aplodontia californica, Grinnell (1913), p. 344.
474 University of Califorma Publications in Zoology | Vou. 17
Type.—A skin and skull in the Berlin Museum.
Type Locality—Assumed to be the Sierra Nevada of California
(Grinnell, 1913, p. 344) ; specimens from Blue Cafion in the central
Sierra may be regarded as typical.
The tenability of Peters’ name for this subspecies depends on
answers to two questions: first, was Peters’ specimen an Aplodontia
at all, and, second, if an Aplodontia, what was its souree? Answering
the first, it seems clear from Peters’ description, which is fairly de-
tailed, which takes account of both skull and skin, and which was
evidently drawn up with Baird’s figure of the skull of Aplodontia
leporina at hand, that the mammal discussed is an aplodontia. The
fact that Peters fell into error in a portion of his description does nos
appear to alter the case from the nomenclatural standpoint. Aplo-
dontia is the only known mammal occurring in California or North
America which possesses approximately the characters set down by
Peters. His references to the inflected angle of the jaw of the animal
he had in hand and to its short tail alone suffice to demonstrate its
position.
The following considerations seem germane to a discussion of the
second question: The discovery of gold in the Sierra Nevada of
California in 1848 brought great numbers of people to this section
from all parts of the world. It is well known that aplodontias occur
in the Sierra not far from the center of early mining activity, and if
is highly probable that some early naturalist collected here the speci-
men which later became the basis of Peters’ new form.*
The type of ‘‘H|[aplodon]. leporinus var. Californicus’’ is stated
‘ ,
to have come ‘‘aus den Gebirgen Californiens.’’ From early times
the Sierra Nevada have impressed travelers and map makers as the
most noteworthy mountains within the state. The only other moun-
tains within its boundaries in which aplodontias occur are the Trinity-
Siskiyou ranges of northern California. The chance that Peters’
specimen came from these then little known mountains rather than
from the Sierra Nevada is slight.
Absolute finality of decision as to the name of the Sierra aplo-
dontia is impossible without access to Peters’ specimen. In deference
1 Soon after the discovery of gold California became comparatively well known,
even in Europe. Phillips (List of Maps of America in the Library of Congress,
1901, pp. 183-186) lists no less than four maps published in Europe between 1848
and 1865 (the date of Peters’ publication of the name californica), according to
which the boundaries of the state are approximately correct. The maps are those
of Duflot de Mofras, Paris, 1849; Desfontaines, Paris, 1849; Reimer, Berlin,
1856; and Rossi, Paris, 1863.
1918 | Taylor: Revision of the Rodent Genus Aplodontia 475
to the above considerations, however, the writer finds himself unable
to agree with Dr. C. Hart Merriam in regarding Peters’ name as
untenable, but is forced to consider that the balance of the evidence
now available favors its validity.
Specimens Examined.—A total of 75, from localities as follows:
California: Siskiyou County—Mt. Shasta, Upper Ash Creek (alt. 7000 ft.),
2; Mt. Shasta, Upper Mud Creek (alt. 7000 ft.), 8. Lassen County—Lassen
Peak, upper edge Canadian zone, east side, 1; Susanville, mountains 12 miles
west, 1. Sierra County—Salmon Lake (alt. 6600 ft.), 5. Placer County—Blue
Cafion, 21. El Dorado County—Lake Tahoe, Emerald Bay, 3; South Fork of
American River, 1. Alpine County—Hope Valley, 10. Mariposa County—Yo
semite Park, East Fork Indian Cafion, 2; near Poreupine Flat, 1; head of Lyell
Canon (alt. 9700 ft.), 5; Mt. Lyell, 4; Chinquapin (alt. 6256 ft.), 5. Mono
County—Mammoth, 4 (2 skulls only).
Geographic Range——The Sierra Nevada of California, from Mt.
Shasta on the north at least to Mammoth, Mono County, on the south.
Zonal range, Boreal.
Cranial Characters.
ilar, in dimensions, to Aplodontia rufa rainierit and A. 1. columbiana,
Skull large (see measurements below), sim-
averaging slightly larger than A. r. rufa; rostrum comparatively long,
zygomatic arches heavy, tending to be squarer anteriorly than in any
other aplodontia; zygomatic arches usually not so much expanded at
the posterior root, viewing cranium on its dorsal or ventral aspect,
as in columbiana; distinct fossa present in typical material on under
surface of expanded portion near posterior root, unlike most examples
of rainieri and columbiana in this respect ; postorbital processes on the
jugal sometimes faintly indicated; temporal ridges tending to ap-
proach to within a few millimeters of each other for their entire length,
variously accentuated in different specimens; caliber of audital tubes
variable, but averaging decidedly greater than in either typical rainiert
or in columbiana, about the same as in rufa; external auditory meatus
round or slightly flattened dorsoventrally, as in rainiert.
External Characters—Above, in summer skins, pale ochraceous-
buff to ochraceous-buff, in most specimens uniformly grizzled with
blackish, and with an insprinkling of silvery tipped hairs; amount and
concentration of black varying to a considerable extent; white spot
at base of ear (pl. 29); underparts light mouse gray to quaker drab,
with insprinkling of black hairs, often silvery tipped, certain speci-
mens with indistinct wash of brown coloration near pinkish buff or
light buff, others with a suggestion of light ochraceous-buff. Color-
ation of winter specimens only slightly different from that in summer.
476 University of California Publications in Zoology | Vou. 17
While the coloration of adults and young is similar, the latter are
woollier and grayer in general appearance than in the former.
Age Variation —Cranially there is a tendency for the sutures be-
tween the nasals posteriorly, the premaxillaries and the frontals to
remain open longer than is the case in other species. The interorbital
constriction undergoes a proportional and with some exceptions an
actual decrease with age. The zygomata are heavier in adults than
in the young and tend to be more bowed outward, or squarer, ante-
riorly. Temporal lines or ridges come closer together with age. In
some old adults of this species they almost coalesce anteriorly, while
remaining only four or five millimeters apart posteriorly. Although
the mastoid process tends to grow laterad more rapidly than the audital
tube with age, in almost all the available specimens the latter exceeds
the former in length.
Sexual Differences—The erania of males tend to be larger than
those of females in nearly all measurements; zygomata are heavier
in males; and the temporal lines or ridges tend to be more accentuated
and more closely approximated. The females tend to have interptery-
goid fossa proportionally broader than the males. In seven out of
eleven males measured the mandible, transversely across angular pro-
cess, is greater than the maximum for the females. Examination of
the material before me confirms C. Hart Merriam’s (1886, pp. 327,
328) conclusions regarding cranial differences due to sex in this species
except in certain details respecting the suture which separates the
frontal bones from the premaxillaries and nasals, and in the outline
of the postzygomatie notches. In Merriam’s material the suture in
question is open in the females and closed in males, while in the
material before me there is nearly as strong a tendency toward efface-
ment of the suture in females as in males. In Merriam’s material
the postzygomatic notches are larger in females, while in the material
now available no constant sexual differences in this respect can be
made out. Our specimens like those examined by Merriam have the
skulls of the females less heavy and massive than those of the males,
the oceipital crest not so highly developed, and the zygomatic arches
not so much bowed outward.
Molt and Seasonal Change.
ing takes place during late summer and early fall. The earliest molt-
ing specimen examined is a female taken July 21. Most examples
taken during August and September are well along in the molting
process, while those secured in October show its last stages, and a
As in other subspecies of rufa, molt-
a
1918 | Taylor: Revision of the Rodent Genus Aplodontia 477
specimen taken on October 15 (no. 192618, ¢, U. S. Nat. Mus., Merriam
Coll.) has the winter pelage complete. An example taken on August
27 (no. 110241, J ad., U.S. Nat. Mus., Biol. Sury. Coll.) shows no sign
of molt, to all appearances having assumed the fresh pelage. One
taken August 29 (no. 110243, 2 ad., U.S. Nat. Mus., Biol. Surv. Coll.)
has the new pelage nearly complete. A few scattered hairs are still
coming in on the sides posteriorly and far back ventrally. These two
specimens have completed the molt at an exceptionally early date.
In one specimen (no. 67854, 2, September 8, U. S. Nat. Mus., Biol.
Surv. Coll.) the pelage has been renewed dorsally on the anterior
three-fourths of the body. In another (no. 192617, ¢, October 12,
U.S. Nat. Mus., Merriam Coll.) all the pelage dorsally has been re-
newed except in a band about two inches wide across the body just
back of the shoulders.
Young specimens do not conform exactly to the schedule of the
adults. In one example (no. 22617, Mus. Vert. Zool.) taken June 19
the new pelage covers the head and extends posteriorly to a line trans-
versely across the body from one to two inches back of the ear; it
appears in a small spot just back of the nape of the neck; and it is
more conspicuous than elsewhere in a broad band about four inches
wide, extending from side to side across the body in the middle of the
back. Often a vigorous hair renewal becomes apparent only upon a
close examination. A juvenal (no. 192615, %, October 9, U. S. Nat.
Mus., Merriam Coll.) in which the hair is being renewed on hips, head,
and neck exemplifies this state of affairs. In a third juvenal (no.
110247, August 30, U. S. Nat. Mus., Biol. Surv. Coll.) new hair is
coming in on areas two inches wide and three inches long far back on
the sides.
The fresh pelage is longer and a trifle browner than the worn
pelage it replaces. Wear has a tendeney to expose the darker under
portions of the pelage, so that the summer pelage appears a little
darker than the fresh fall pelage. The differences between the pelages
are slight at most, and often none are observable.
Remarks.—A plodontia rufa californica differs from A. r. rufa of
the Trinity-Siskiyou region of southern Oregon and northern Cali-
fornia in having more grayish coloration and in cranial characters as
follows: Nasal outline uniform in californica, moderately broad in
front, moderately narrow posteriorly (nasals in rufa variable, usually
proportionally broader anteriorly and narrower posteriorly) ; widest
portion of nasals usually most anterior pomt where nasals touch
478 University of California Publications in Zoology — [Vou. 17
maxillaries (often posterior of this point in rufa); shallow embay-
ment in lateral outline of nasals posteriorly (variable in rufa) ; ros-
trum and nasais longer in californica, zygomatic arches squarer ante-
riorly, caliber of audital tubes averaging greater (about equal to eali-
ber in typical rufa). From A. r. rainieri, californica differs in the
following skull characters: nasals tending to be broadest at a more
anterior point; a more pronounced tendency observable toward ap-
proximation of temporal ridges; heavier zygomatic arches, squarer
anteriorly ; and great caliber of audital tubes.
CRANIAL MEASUREMENTS OF Aplodontia rufa californica (14 SKULLS)
Average Mean Maximum Minimum
Bastar Wen gts cccyesc ese sseece Necro eee 61.1 62.4 65.1 59.6
Length of nasals (8 skulls) 26.8 26.9 28.4 25.4
Widthvot masals, -- see iei 12.0 12.8 11.2
Length of audital tube (12 skulls) -.. 19.1 18.9 20.5 WEP)
Length of incisive foramen ... 7.6 7.6 8.2 7.1
Zygomatic width .......2...-...-... 57.6 57.0 60.9 53.1
Greatest width of interpterygoid fossa 5.2 5.2 5.8 Awl
Mastoid width of cranium .............. 54.2 53.9 57.0 50.7
Alveolar length of superior cheek teeth 19.6 19.6 20.7 17.5
Distance between infraorbital foramina .-.... = 16 17.4 18.4 16.4
Mandible, transversely across angular process ... 24.5 24.4 26.6 22.1
Mandible} greatest length) [= ee 50.5 49.9 53.2 46.7
EXTERNAL MEASUREMENTS OF Aplodontia rufa californica (17 SKINS)
Average Mean Maximum Minimum
Total length 352 352 380 325
JERUOCL BHO OLE se coreerpeceteeecceuetore ce 60 59 64 Bis)
Cranial measurements of three males from Blue Cafion, California: basilar
length, 64.3, 63.1, 62.8; length of nasals, 29.1, 27.0, 27.0; width of nasals, 12.6,
13.0, 12.8; zygomatic width, 61.9, 61.4, 61.2; mastoid width, 59.4, 59.9, 57.9.
Cranial measurements of three females from the same locality: basilar length,
60.7, 61.9, 60.5; length of nasals, 27.6, 27.2, 27.0; width of nasals, 12.0, 11.9, 12.2;
zygomatie width, 55.5, 60.9, 57.8; mastoid width, 53.4, 57.9, 54.8.
Californica is one of the more consistent subspecies of Aplodontia
rufa, maintaining its characters with considerable uniformity through-
out its range. Still, a series of specimens from Mount Shasta, Cali-
fornia, while indistinguishable cranially from typical californica, is
characterized by slightly darker coloration dorsally.
Squareness anteriorly of the zygomatic arches and outline of the
nasals in specimens of rufa from Mount Mazama in southern Oregon
suggest intergradation with californica, as do also certain cranial
characteristics of examples from Canon Creek, California. The latter
exemplify a tendency to combine the square zygomatic arches and the
distinctive nasal outline of californica with the short rostrum and
browner coloration of rufa.
1918] Taylor: Revision of the Rodent Genus Aplodontia 479
Aplodontia rufa nigra Taylor
Point Arena Aplodontia
Aplodontia nigra Taylor (1914), pp. 297-300.
Type—Male adult; no. 20320, Mus. Vert. Zool.; Point Arena,
Mendocino County, California; July 10, 1913; collected by ©. L.
Camp; orig. no. 1003.
Specimens Examined.—Total number 4, all from California: Men-
docino County—Point Arena.
Geographic Range-—Known only from the type locality, where it
is found within an area of approximately twenty-four square miles.
Cramal Characters—Skull moderate in size (see measurements
below) ; nasals dilated anteriorly, comparatively contracted poste-
riorly ; zygomatic arch medium in weight, squarish anteriorly ; position
of postorbital process faintly indicated ; temporal lines or ridges not
closely approaching ;. incisive foramina short; noteh dorsally on ex-
ternal auditory meatus deeper than in Aplodontia rufa humboldtiana,
more as in A. r. pacifica.
External Characters—Summer specimens, above shiny black, with
a sparse insprinkling of pinkish buff hairs and with plumbeous bases
of the hairs showing through to a certain extent, the whole giving the
impression of shiny black faintly sprinkled with grayish; sides paler
than back, prevailingly pinkish buff, with heavy insprinkling of black
hairs; head tending to be shiny black; face dark quaker drab; under-
parts cinereous to plumbeous, lightly washed with pinkish buff.
Age Variation—This subspecies exhibits a remarkable similarity
in the characters of adult and young. The coal black dorsal color-
ation, as well as the anterior dilation of the nasal outline, are conspic-
uous in both. In one young example (no. 20321, Mus. Vert. Zool.)
a single small bony element is marked off by sutures in the inter-
parietal region. The zygomata are decidedly heavier in the adult than
in the young, as well as somewhat more expanded anteriorly.
Molt—Three young at hand (nos. 20318, 20319, and 20321, Mus.
Vert. Zool., taken July 9 to 11) are molting from the juvenal pelage
into that of the adult. The new pelage is more intensely brown and
black than the gray-black pelage of the juvenal, the brown being more
emphasized laterally, the black dorsally.
Remarks —Aplodontia rufa nigra is the most strikingly marked
subspecies of mountain beaver known, its dorsal coloration rendering
480 University of Califorma Publications in Zoology — [Vou. 17
it the darkest form as yet characterized. Both old and young may
be separated with certainty on coloration alone from any known form
of the genus. The anterior dilation of its nasal outline serves to sep-
arate it from A. r. phaea and A. r. pacifica. From A. r. humboldtiana,
its nearest neighbor on the north, it is separated by its darker color-
ation and, in general, smaller size.
CRANIAL MEASUREMENTS OF Aplodontia rufa nigra (1 CRANIUM)
Basilar: len oth’ ic. 82 ics acsncsaten anes se oe eae eee eee 57.1
Tengphion nasalls' ts: e ee ee ene ee 23.2
Width of nasals ......... 11.0
Length of audital tube ....... aU
Length of incisive foramen
ZY FOMAtIC wpb) Severe eeceesee ees eee
Greatest width of interpterygoid fossa
Mastoid width of cranium .......................
Alveolar length superior cheek teeth -.
Distance between infraorbital foramina _..
Mandible, transversely across angular process .....
Mandible;yoreatest lengthy =. ere ee
oO
OS) Me ol ed oor)
Paw ODN OW
HB DORR OF
EXTERNAL MEASUREMENTS OF Aplodontia rufa nigra (1 SK)
Motallen'gbhies wesc 346
PLN hy POO 1 2 Seo sack sence cen eee see eons 55
That Aplodontia rufa nigra finds in A. r. humboldtiana and in
A. r. phaea its nearest relatives seems reasonably clear. The blackish
coloration of the Point Arena form suggests the dark coloration of
the Humboldt Bay race, while its size and the length of incisive fora-
mina of the skull show that it is not far from phaea.
The distinctive coloration of all the known specimens of A. r. nigra
and its geographical isolation constitute, perhaps, arguments for its
recognition as a full species. On the other hand, the paucity of
material representative of the form, coupled with the amplitude of
individual variation in other forms in the genus, and with the evident
overlapping of cranial characters with humboldtiana on the one side
and phaea on the other, indicate the propriety, for the present at least,
of allocating 2igra as a subspecies of rufa.
Aplodontia rufa phaea Merriam
Point Reyes Aplodontia
?Haplodon rufus, True (1885), p. 596.
Aplodontia phaea Merriam (1899a), p. 20.
Haplodontia phaea, Elliot (1901), p. 114.
Type.—Male adult, skin and skull; no. 186475, U. S. Nat. Mus.,
Merriam Coll. (no. {)); Point Reyes, Marin County, California;
August 1, 1886; collected by C. A. Allen; orig. no. 142.
1918 | Taylor: Revision of the Rodent Genus Anlodontia 481
Specimens Examined.—A total of 36, from the following localities:
California: Marin County—Five miles west of Inverness, 9; six miles west of
Inverness, 7; four miles south of Olema, 3; Lagunitas, 2; Point Reyes, 15 (1, skull
only; 3, jaws only, labeled under one number).
Geographic Range—FKavorable situations in Marin County, Cali-
fornia, where it is found within an area of approximately 110 square
miles.
Cranial Characters.—Skull small (see measurements below) ; nasals
not extreme, nearly straight sided, narrow across posterior ends, not
extremely wide at anterior ends; zygomatic arch light, not conspicu-
ously squarish anteriorly ; temporal lines or ridges not unduly accent-
uated, wide apart; audital tube comparatively large calibered ; fossae
anterior of lambdoidal crests shallow ; notch dorsally on external audi-
tory meatus shallow; incisive foramina short; interpterygoid fossa
narrow.
External Characters Above, 1 summer skins, pinkish cinnamon
to cinnamon-buff, in winter approximating cinnamon or even tending
toward sayal brown, sometimes grayish; all upperparts uniformly
grizzled with black-tipped hairs, which are somewhat more numerous
on the back than on the sides. Underparts French gray to plumbeous,
with a sparse insprinkling of black hairs; the whole ventral surface
washed with light ochraceous-buff to pinkish buff. Two specimens
(nos. 192629, 192631, U. S. Nat. Mus., Merriam Coll.) have small white
spots on the throat. One example (no. 192635, U. 8. Nat. Mus., Mer-
riam Coll.) is melanistic, being light seal brown in color above and
below.
Age Variation.—Ilustrative of the decrease in width of the inter-
orbital constriction with age are the measurements obtained from ten
specimens of this species, in which the ratio of the width of inter-
orbital constriction to basilar length varies from 26.2 per cent in a
very young example to only 17.1 per cent in an adult.
In several of the youngest specimens there are two small bones
marked off by sutures in the interparietal region. They soon dis-
appear through ankylosis.
In ventral view it may be noted that with age the part of the palate
between the third premolar and the incisive foramina becomes nar-
rower. One striking feature noticed here is that while the basilar
length varies with age in certain specimens examined from 42.9 mm.
to 55.5 mm., the width of the palate between right and left premolar
four remains constant at about 5 millimeters.
482 University of Califorma Publications in Zoology [ Vou. 17
The ratio of the width of the angular portion of the jaw to the
length of the jaw undergoes, in specimens measured, a 5 per cent
increase with age. In youth there is a smooth space between the inner
prominence of the angle and the ridge in front of the masseteric fossa,
while in old age there is a tendency for this ridge to be continued
Figs. N-P. Dorsal view of crania of young specimens of Aplodontia rufa
phaea, to illustrate marking off of interparietal elements. X 1. Fig. N, no.
20302, Mus. Vert. Zool.; fig. O, no. 20308, Mus. Vert. Zool.; fig. P, no. 20316,
Mus. Vert. Zool.
1918 | Taylor: Revision of the Rodent Genus Aplodontia 483
across the lower side of the ramus and connected with the inner promi-
nence of the angle.
Seasonal Change——Molt begins the last of July or the first of
August. Specimens taken September 17 and November 9 have the
process about completed, and one secured November 12 has entirely
finished molting. More black hairs are sprinkled into the winter
pelage, and the hair is somewhat longer and thicker. The spot marks
about the mammae in the females are much less conspicuous in winter
than in summer.
CRANIAL MEASUREMENTS OF Aplodontia rufa phaea (11 SKULLS)
Average Mean Maximum Minimum
IBS AS Ma PLO OG ip sea cece ec Sos ecto cost ezcceesececeece -- 59.6 54.8 58.6 51.1
Length of nasals (4 skulls) ... 21.5 22.0 23.7 20.3
Width) of masals) -4.2-..-<--..2.2.-: 9.6 9.5 10.3 8.7
Length of audital tube (9 skulls) 16.3 16.4 17.2 15.6
Length of incisive foramen 6.2 6.1 7.6 4.5
Zygomatie width .................----- 50.3 49.5 53.0 46.0
Greatest width of interpterygoid fo : - 43 4.2 4.7 3.8
Mastoid width-of cramium ..-........-2--<-:--cc-cecscse0--- 47.2 46.4 51.0 41.9
Alveolar length superior cheek teeth 0... 18.4 18.5 19.2 iW)
Distance between infraorbital foramina ney el De 15.3 16.5 14.1
Mandible, transversely across angular Pee pee 2186 21.7 22.8 20.7
Mandible, PTeATe Salen Oi Ne ecekssexsevecsce tees ance renee 43.1 43.4 45.1 eeley/
EXTERNAL MEASUREMENTS OF Aplodontia rufa phaea (13 SKINS)
Average Mean Maximum Minimum
308 312 344 280
52 52 57 48
Total length ..
Hind foot ....
Remarks.—Aplodontia rufa phaea is the smallest subspecies in the
genus. It is also the palest of the coastal forms of Aplodontia (pa-
ctfica, humboldtiana, nigra, phaea) occurring in Oregon and California.
Cranially its closest affinities seem to be with Aplodontia rufa nigra.
In spite of the geographical isolation of the Marin County form,
it is very similar, in general coloration, to some examples of rufa; nor
are its cranial characters sufficiently distinctive to warrant its recog-
nition as of more than subspecifie rank.
484 University of California Publications in Zoology | Vou. 17
F. LITERATURE CITED, WITH TITLES OF OTHER WORKS
CONTAINING MATTER ON APLODONTIA
ALLEN, J. A.
1892. Visitors guide to the collection of mammals in the American Museum
of Natural History. (New York, Amer. Mus. Nat. Hist.), 89 pp.,
24 figs. in text.
Sewellels or showtls, family Aplodontidae, mentioned (p. 31).
ALSTON, E. R.
1876. On the classification of the order Glires. Proce. Zool. Soe. London,
1876, 61-98, pl. 4, 5 figs. in text.
Haploodontidae placed with Sciuromorpha (pl. 4, opp. p. 61);
family name spelled Haplodontidae (pp. 66, 75, 78); generic name
Haplodon used (p. 78); outline of characters of family given
(p. 78).
AntTuHony, H. E.
1916. Habits of Aplodontia. Bull. Amer. Mus. Nat. Hist., 35, 53-63, 8 figs.
Habits of Aplodontia pacifica as observed in vicinity of Tilla-
mook, Oregon.
AUDUBON, J. J., and BACHMAN, J.
1854. The quadrupeds of North America. (New York, Audubon), 3, vi+348,
pls. 101-155.
Aplodontia leporina diseussed (pp. 99-103); figured (pl. 123).
Bairp, 8S. F.
1857. Report on Zoology. I. Mammals. U.S. Pac. R. R. Expl. and Surv.,
8, xlviii + 757, 35 figs. in text. pls. 17-60 (except pl. 29, published
elsewhere).
Classification, description and history of genus Aplodontia dis-
cussed (pp. 350-352); descriptive remarks and comments on Aplo-
dontia leporina (pp. 353-354).
BuytH, E.
1840. Mammalia, in Cuvier’s Animal Kingdom. (London, Orr), pp. 38-152,
figs. 2-66 in text.
Name Aplodontia used for author’s ‘‘abnormal’’ phytophagous
order of placental mammals (pp. 150, 152).
Bonaparte, C. L.
1831. Saggio di una distribuzione metodica degli animali vertebrati. Gior-
nale Arcadico di Scienze Lettere ed Arti, 49, 1-77. (This separate
repaged; pagination incorrect, ie., p. 5 of separate is p. 4 of
original, ete.)
Genus Aplodontia listed (p. 20).
Branpt, J. F.
1855. Beitriige zur nihern Kenntniss der Siiugethiere Russlands. Mem.
Acad. Sci. Nat. St. Petersbourg, ser. 6, 7, 1-365, 17 tables.
Spelling of name of genus Haploodon; also relationships of the
genus discussed (pp. 150-151).
1918 | Taylor: Revision of the Rodent Genus Aplodontia 485
BRETHERTON, B. J.
1895. Some Oregon mammals: The mountain boomer (Haplodon rufus).
Oreg. Nat., 2, 123-125, 1 fig. in text.
Discussion of habits.
Brooks, A.
1899. The sewellel, Aplodontia rufa. Recreation, 2, 258-259, 1 halftone.
Habits as observed in southern British Columbia.
Bryant, W. E.
1891. A provisional list of the land mammals of California. Zoe, 1, 353-360.
Aplodontia major listed (p. 355).
1892. Recent additions to the North American land mammal fauna. Zoe,
3, 201-223.
Aplodontia Richardson is said to antedate Haplodon Richardson
[?] (p. 203).
CALIFORNIA ACADEMY OF NATURAL SCIENCES,
1855. [Meeting for September 24, 1855], Proc. Calif. Acad. Nat. Sci., 1, 71.
Record of donation to the cabinet of a ‘‘species of marmot,
perhaps undescribed’’ from the vicinity of the Great Trees, Cala-
veras County. ‘‘The miners call it Mammoth Mole.’’
1866. [Meeting for September 18, 1865], Proc. Calif. Acad. Nat. Sci., 3, 224.
Record of donation of a ‘‘Specimen of Aplodontia leporina, shot
near Lake Tahoe, by J. M. M’Donald.’’
Cassin, J.
1858. Mammalogy and ornithology. U.S. Expl. Exp. (Wilkes), 8, viii+466,
several unnumbered figures in text.
Compilation of matter on Aplodontia leporina; quotes extensively
from Peale’s vol. 8 of the same series; skull figured (pp. 36-37).
Cooper, J. G.
1860. Report upon the mammals collected on the Survey, U. 8S. Pac. R. R.
Expl. and Surv., 12, Zool. Rep., bk. 2, pt. 3, no. 2, chap. 1, pp. 73-88.
Distribution and habits of Aplodontia leporina (p. 82).
1868. Zoology, in Cronise, The natural wealth of California. (San Francisco,
Bancroft), pp. 434-501. ;
Reference to Aplodontia leporina, with several lines of comment
regarding its habits and status in California (p. 442).
Corr, E. D. :
1878. On some characters of the Miocene fauna of Oregon. Palae. Bull.,
30, 1-16.
Original deseription of Meniscomys hippodus, new genus and
species (pp. 5-6); and of Meniscomys multiplicatus, new species
(p. 6).
1883. The extinct rodentia of North America. Amer. Nat., 17, 438-57, 13
figs. in text. :
Haplodontiidae mentioned in comparisons with Castoridae and
Sciuridae (p. 54); reference to Haplodonta [= Haplodontia] rufa
(p. 55).
CouEs, E.
1877a. The mountain boomer, or showtl. Amer. Nat., 11, p. 434.
Paragraph introductory to publication of letter from Dr. F. 8.
Matteson regarding habits of Aplodontia leporina.
486 Unwersity of California Publications in Zoology | Vou. 17
1877b. ‘‘Haplodontidae’’ in Monographs of North American Rodentia, in
Rep. U. 8. Geol. Surv. Terr., 11, no. 9, pp. 543-601, 1 pl.
Discussion of characters and relationships of family, genus and
species, including remarks on history, relationships and habits;
accompanied by one plate (pl. 6) illustrating dorsal, lateral and
ventral view of skull (pp. 549-599).
1890. Haplodon. Century Dictionary (New York, Century), Part x, p. 2712.
Discussion of spelling of the generic name for Aplodontia.
DeEsMAREST, M. A. G.
1822. Mammalogie, ou description des espéces de mammiféres. (Paris,
Agasse), pp. 1-viii, 277-530. The whole = vol. 126 of the Encyclo-
pédie methodique.
Unimportant fragmentary compilation of matter on ‘‘ Anisonyx?
rousse’’ (footnote, p. 330).
Doue.as, D.
1836a. A sketch of a journey to the northwestern parts of the continent of
North America, during the years 1824-1827, in Hooker, W. J.,
Companion to Botanical Magazine (London, Curtis), 2, 83-140.
The Ground Rat, or Arctomys (Arctomys brachyurus?), mentioned
as occurring on the Cowalidsk River (p. 101).
1836b. Account of Mr. Douglas’ second visit to the Columbia; his excursions
in California; and his visit to Mouna Roa in the Sandwich Islands;
with particulars respecting his death. Ibid., 2, 146-178.
Remarks on geographic limits of California (p. 149).
1914. Journal kept by David Douglas during his travels in North America,
1823-27 (London, Wesley), 364 pp., frontispiece.
Arctomys [= Aplodontia|] mentioned (pp. 59, 156); ground rat
(p. 156).
Exxior, D. G.
1899. Catalogue of mammals from the Olympic Mountains, Washington,
with descriptions of new species. Field Columb. Mus. Zool., 1,
241-276, pls. 41-61, several unnumbered figs. in text.
Treatment of Haplodontia olympica, with illustrations, and casual
mention of H. rufa and H. major (pp. 251-253, pls. 41, 42).
1901. A synopsis of the mammals of North America and the adjacent seas.
Field Columb. Mus. Zool., 2, xiv+471, 49 pls., 94 figs. in text.
Haplodontidae characterized (p. 111); six forms of Haplodontia
listed, as follows: Haplodontia rufa; H. r. californica; H. r. rainieri;
H. pacifica; H. phaca; H. olympica (pp. 112-114).
1903. A list of mammals obtained by Edmund Heller, collector for the
Museum, from the coast region of northern California and Oregon.
Field Columb. Mus. Zool., 3, 175-198.
References to Haplodontia (pp. 175, 179, 184, 185); Haplodontia
pacifica (p. 184); H. phaea (pp. 184-185).
1905. A check list of mammals of the North American continent, the West
Indies and the neighboring seas. Field Columb. Mus. Zool., 6,
v+761, frontispiece.
Genus Haplodontia listed (p. 126); six forms of Haplodontia
enumerated, as follows: Haplodontia rufa; H. r. californica; H. r.
rainieri; H. pacifica; H. phaea; H. olympica (pp. 126-128).
1918] Taylor: Revision of the Rodent Genus Aplodontia 487
1907. A catalogue of the collection of mammals in the Field Columbian
Museum. Field Columb. Mus. Zool., 8, vili+694, 92 figs. in text.
Four forms of Haplodontia listed as being represented in the
Museum Collection, as follows: Haplodontia rufa; H. pacifica; H.
phaea; H. olympica (pp. 188-190).
FiscuHer, J. B.
1830. Synopsis mammalium. (Stuttgart, Cotta), xli + 752 pp.
Comments given on Apludontia leporina; of little importance
(addenda, pp. 330, 398, 399, errors for 530, 598, 599).
ForsytH Magor, C. J.
1893. On some Miocene squirrels, with remarks on the dentition and elassi-
fication of the Sciurinae. Proc. Zool. Soc. London, 1893, 179-215,
pls: 8—lil:
Refers to Winge’s classification of rodents. Sciuridae and
Anomaluridae derived from the Haplodontidae (p. 196, footnote).
Furtone, E. L.
1910. An aplodont rodent from the Tertiary of Nevada. Univ. Calif. Publ.
Bull. Dept. Geol., 10, 397-403, 6 figs. in text.
Original description of Aplodontia alexandrae.
GEOFFROY SaINt-Hiarrg, I.
1826. La marmotte rousse, Arctomys rufa. Dict. Class. d’Hist. Nat. (Paris
Baudouin Fréres), 10, 1-642.
General compiled account of Aplodontia, headed as above; also
discussion of old world and new world species referred to Arctomys
(pp. 186-187).
)
GERRARD, FE.
1862. Catalogue of the bones of mammalia in the collection of the British
Museum (London, British Museum), iv-+296 pp.
A skull of Aplodontia leporina from Chilakiveyak [= Chilli-
wack] River, British Columbia (p. 224), and another from the
Wilkes Exploring Expedition (addenda, p. 296) are in the collee-
tion.
GERVAIS, P.
1854. Histoire naturalle des mammiféres. (Paris, Curmer), xxiv + 420 pp.,
pls. 19, unnumbered figs. in text. The ‘‘Genre Aplodontie’’ refer-
red to the ‘‘Tribu des Saccophoriens’’ of the ‘‘Famille des Pseu-
dostomides’’; associated with the genus Saccophorus (p. 364).
GIEBEL, C. G.
1855. Die Saugethiere. (Leipzig, Abel), xii + 1108.
Haplodon referred to the family Spalacini; the genus Haplodon
(p. 526); H[aplodon]. leporinus (p. 527).
1859. Die Naturgeschichte des Thierreichs. JI. Die Siiugethiere. (Leip-
zig, Wigand), viii + 522 pp., 926 figs. in text.
Reference made to Haplodon leporinus in account of genus
Georhychus ; skull figured (p. 279).
Ginn, T.
1872. Arrangement of the families of mammals with analytical tables.
Smithson. Mise. Coll., 11, vi + 98.
Haploodontidae listed as sole family under superfamily Hap-
loodontoidea (p. 22).
488 University of California Publications in Zoology (Vou. 17
Gray, J. E.
1843. List of the specimens of mammalia in the collection of the British
Museum. (London, British Museum), xxviii + 216 pp.
Three skins of the Sewellel, Aplodontia leporina, from ‘‘N.
America,’’ are in the collection (p. 150).
GRIFFITH, E.
1827. The animal kingdom. (London, Whittaker), 5, 1-391.
Reference to A[retomys]. rufa (p. 245).
GRINNELL, J.
1913. A distributional list of the mammals of California. Proce. Calif. Acad.
Sci. (4), 3, 265-390, pls. 15, 16.
Two species of Aplodontia—aA. californica and A. phaea—listed
as occurring in California (p. 344).
1915. The vertebrate fauna of the Pacifie Coast, in Nature and science on
the Pacific Coast. (San Francisco, Elder), pp. 104-114, pls. 12-14.
Aplodontia mentioned (p. 110); photograph of live animal (pl.
14, opp. p. 118).
1916. An analysis of the vertebrate fauna of the Trinity region of northern
California. Univ. Calif. Publ. Zool., 12, 399-410.
In course of analysis mentions A[plodontia]. californica (p. 401),
and Aplodontia chryseola (pp. 401, 402, 407).
Haran, R.
1825. Fauna Americana. (Philadelphia, Finley), pp. i-x, 11-318.
Compilation of matter on Arctomys rufa (addenda, pp. 308—
309).
Hover, J. B.
1877. History of the American Fauna, in the Museum of Natural History.
(New York, Virtue), pp. i-cexc, 11 pls.
Short compilation; treats family Haplodontidae, genus Hap-
lodon and the species Haplodon rufus (p. xc).
KeEtoee, L.
1910. Rodent fauna of the late Tertiary beds at Virgin Valley and Thou-
sand Creek, Nevada. Univ. Calif. Publ. Bull. Dept. Geol., 5, 421—
437, 20 figs. in text.
Aplodontia alexandrae briefly discussed (p. 429).
1912. Pleistocene rodents of California. Univ. Calif. Publ. Bull. Dept.
Geol., 7, 151-168, 16 figs. in text.
Discussion of Aplodontia major fossilis from Samwel and Potter
Creek caves in California (pp. 157-158); incidental mention of
Aplodontia major (p. 158).
1914. Aplodontia chryseola, a new mountain beaver from the Trinity region
of northern California. Univ. Calif. Publ. Zool., 12, 295-296.
Original description of Aplodontia chryseola.
1916. Report upon mammals and birds found in portions of Trinity, Siskiyou
and Shasta counties, California. Univ. Calif. Publ. Zool., 12,
335-398, pls. 15-18, 1 fig. in text.
Relationships, occurrence and habits of Aplodontia chryscola
(pp. 369-372).
1918] Taylor: Revision of the Rodent Genus Aplodontia 489
Lantz, D. E.
1917. Destroying rodent pests on the farm. Yearbook, U. 8. Dept. Agr.,
Separate 708 (1916), 1-18, 5 pls., 1 fig. in text.
Damage done by aplodontia to crops in western Washington
mentioned (p. 16); half-tone of aplodontia (plate 1, fig. 1).
Lesson, R. P.
1827. Manuel de mammalogie ou histoire naturelle des mammiféres( Paris,
Roret), xv + 442 pp.
Short paragraph on Anisonyx roux (p. 240); evidently com-
piled from Rafinesque and Harlan.
LEwIis, M., and CLark, W.
1814. History of the Lewis and Clark Expedition. (Paul Allen edition,
Philadelphia, Bradford), 2, ix + 522.
Description of ‘‘Sewellel’’ (pp. 176-177).
1876. An account of the various publications relating to the travels of
Lewis and Clark, with a commentary on the zoological results of
their Expedition; Elliott Coues, in U. S. Geol. and Geog. Surv.
Terr. (2), Bull. 6, 417-444.
Short compilation regarding the Sewellel (p. 437).
1893. History of the Expedition under the command of Lewis and Clark.
4 vols. (Coues edition, New York, Harper), 3, i-vi, 821-1298.
Extract on the Sewellel from the journals of Lewis and Clark
(pp. 861-862); critical remarks (footnote, p. 861).
1904-5. Original journals of the Lewis and Clark Expedition, 1804-1806.
8 vols. (Thwaites’ edition, New York, Dodd), 8 [Atlas], i-xvi,
54 maps.
Parts 1 and 2 of map 32 are of particular interest as showing
the localities on the Columbia River visited by Lewis and Clark.
LILLJEBORG, W.
1866. Systematisk ofversigt af de gnagande Daggdjuren, Glires. (Upsala,
Kongl. Akad. Boktryckeriet), pp. 1-59. :
Family Haploodontidae placed between Sciuridae and Chinchil-
lidae in table, opp. p. 9; family discussed (p. 41).
Lorp, J. K. i
1866. The naturalist in Vancouver Island and British Columbia. 2 vols.
(London, Bentley), 1, xiv + 358, frontisp., 6 illus. in text.
Chapter 13 devoted to discussion of habits, Indian names, and
distribution of Aplodontia leporina.
Lum, 8. K.
1878. The sewellel or show’tl. Amer. Nat., 12, January, 10-13.
Remarks on distribution and habits of Haplodon rufus.
[ LyDEKKER, R. |
1914. Guide to the galleries of Mammals in the Department of Zoology of
the British Museum (Natural History), (ed. 9, London, British
Museum), 123 pp., 1 pl. 3 plans, 65 figs. in text.
Reference to Aplodontiidae and the genus Aplodontia (p. 57).
Lyon, M. W., Jr.
1907. Notes on mammals collected at Mount Rainier, Washington. Smith-
sonian Mise. Coll., 50, 89-92.
Aplodontia major rainieri discussed (p. 91).
490 Umversity of California Publications in Zoology — | Vou. 17
Lyon, M. W., Jr., and Oseoop, W. H.
1909. Catalogue of the type-specimens of mammals in the U. 8. National
Museum, including the Biological Survey collection. U. S. Nat.
Mus., Bull. 62, x + 325.
The types of three forms of Aplodontia were in the Biological
Survey Collection on the date of publication of this catalogue, as
follows: <Aplodontia olympica, A. pacifica, and A. major rainieri
(pp. 159-160).
Marsy, O. C.
1877. New vertebrate fossils. Amer. Jour. Sei. (3), 14, 249-256.
Original description of the genus Allomys (p. 253).
Marreson, F. S.
1877. The mountain boomer, or showtl. Amer. Nat., 11, 434-435.
Letter regarding habits of aplodontia.
MarruHew, W. D., and Giptey, J. W.
1904. New or little known mammals from the Miocene of South Dakota.
Amer. Mus. Exp. 1903. Bull. Amer. Mus. Nat. Hist., 20, 241-268,
15 figs. in text.
Meniscomys sp. indt., recovered in Rosebud beds; comparison of
Meniscomys with Haplodontia (pp. 263, 264).
MarrHew, W. D.
1907. A Lower Miocene fauna from South Dakota. Bull. Amer. Mus. Nat.
Hist., 23, 169-219, 26 figs. in text.
Meniscomys sp. listed from Lower Rosebud (p. 172).
1910. On the osteology and relationships of Paramys, and the affinities of
the Ischyromyidae. Bull. Amer. Mus. Nat. Hist., 38, 43-72, 19 figs.
in text.
Aplodontia and the Aplodontiidae considered in comparison and
treatment of primitive rodents (pp. 44, 47, 48, 64-69, 71).
MerRIAM, ©. H.
1886. Description of a new species of Aplodontia (Aplodontia major, sp.
nov.) from California. Ann. N. Y. Acad. Sci., 3, no. 10, pp. 312-
328, 2 pls., 1 fig. in text.
Description of Aplodontia major, including discussion of history,
nomenclature, habits, and variation.
1897. The mammals of Mount Mazama, Oregon. Mazama, 1, 204-230.
Habits of Aplodontia major as observed on Mount Mazama,
near Crater Lake (p. 219).
1899a. Descriptions of six new rodents of the genera <Aplodontia and
Thomomys. Proc. Biol. Soe. Wash., 18, 19-21.
Original descriptions of Aplodontia pacifica, A. phaea, A. olym-
pica, and A. major rainieri.
1899b. Results of a biological survey of Mount Shasta, California. U. 8.
Dept. Agr., Div. Biol. Surv., N. Amer. Fauna, 16, 1-179, pls. 1-5,
46 figs. in text.
Aplodontia major rainieri listed as a Cascade species (pp. 74,
81); occurrence and habits of Aplodontia major on Mt. Shasta
(pp. 92, 93).
1918] Taylor: Revision of the Rodent Genus Aplodontia 491
Merriam, J. C.
1911. Tertiary mammal beds of Virgin Valley and Thousand Creek in
northwestern Nevada. Part II, Vertebrate faunas. Univ. Calif.
Publ. Bull. Dept. Geol., 6, 199-304, pls. 32-33, 80 figs. in text.
Aplodontia alexandrae listed from Virgin Valley (p. 205); from
Thousand Creek (p. 211).
1916. Tertiary vertebrate fauna from the Cedar Mountain region of west-
ern Nevada. Univ. Calif. Publ. Bull. Dept. Geol., 9, 161-198, pl.
8, 48 figs. in text.
Aplodontia sp. from the Upper Miocene described (pp. 177-179).
Merriam, J. C., and Sinciair, W. J.
1907. Tertiary faunas of the John Day region. Univ. Calif. Publ. Bull.
Dept. Geol., 5, 171-205.
Four species of Allomys from the John Day beds are assigned
to the Haplodontidae (p. 185).
Minter, G. S., Jr.
1912. List of North American land mammals in the U. 8S. National Museum,
1911. U.S. Nat. Mus., Bull. 79, xiv + 455.
Six forms of Aplodontia listed, as follows: <Aplodontia major
major, A. m. rainieri, A. olympica, A. pacifica, A. phaea, A. rufa
(p. 291).
Miter, G. S., Jr., and REN, J. A. G.
1901. Systematic results of the study of North American land mammals to
the close of the year 1900. Proe. Bost. Soc. Nat. Hist., 30, 1-352.
Six forms of Aplodontia listed, as follows: Aplodontia major,
A. m. rainieri, A. olympica, A. pacifica, A. phaea, A. rufa (pp. 63-64).
MINDING, J.
1829. Ueber die geographische Vertheilung der Siiugethiere. (Berlin, Enslin),
pp. 1-104.
Refers genus Anisonia [= Anisonyx— Aplodontia (part)]| to
family Prensiculantia; genus includes species brachyura and
Smiubpac 2 \(p- 86).
Muir, J.
1909. Our national parks. (Boston, Houghton), x + 382 pp., numerous un-
numbered plates.
Reference to Haplodon (p. 201).
Murpny, J. M.
1876. The hunting fields of the Pacific Coast. Capturing the showtl. Rod
and Gun, May 20, p. 121.
Habits of Aplodontia leporina.
1877. The shewelel or showtl. London Field, April 28, 486-487.
Habits of Aplodontia leporina.
Murray, A.
1866. The geographical distribution of mammals (London, Murray), xvi +
420 pp., 101 maps, numerous figs in text, numbered separately under
chapters.
Reference to classification and habits of Aplodontia leporina
(p. 263); additional references (pp. 356, 399, 401).
492 University of California Publications in Zoology — (Vou. 17
NeEwserry, J. S.
1857. Report upon the mammals, in Zoological Report, in U. S. Pac. R. R.
Expl. and Surv., 6, 35-72, 3 pls.
Remarks on distribution of Aplodontia leporina (p. 58).
Osporn, H. F.
1910. The age of mammals in Europe, Asia and North America. (New
York, Macmillan), xvii + 635 pp., 220 figs. in text.
Superfamily Aplodontoidea; also family Aplodontiidae with
three genera, Meniscomys, Mylagaulodon, Aplodontia, listed (p.
534).
PauLMER, T. S.
1904. Index generum mammalium: a list of the genera and families of
mammals. U.S. Dept. Agr., Div. Biol. Surv., N. Amer. Fauna, 23,
pp. 984.
Possible variations in generic name Aplodontia discussed (p.
25); genus Aplodontia (pp. 112-113); family names considered
(pp. 744, 782, 841-842).
Pras, I. RR:
1848. Mammalia and ornithology. U. S. Expl. Exp. (Wilkes) (Phila-
delphia, Sherman), 8, i-xxvi, 17-338, several unnumbered figs. in
text.
Aplodontia leporina discussed, and skull figured (pp. 56-57).
PETERS, W.
1864. Neue Arten der Saugethiergattungen Geomys, Haplodon, und Dasypus.
Mon. k. preuss. Akad. Wiss. Berlin, 1864, 177-180.
Original description of H[aplodon]. leporinus var. californicus.
PRICE, W. W.
1894. Notes on a collection of mammals from the Sierra Nevada Mountains.
Zoe, 4, 315-332.
Habits and distribution of Aplodontia major (p. 328).
RAFINESQUE, C. S.
1817. Descriptions of seven new genera of North American quadrupeds, in
Museum of Natural Sciences, in Amer. Monthly Mag. and Crit.
Rev., 2, 44-46.
Original description of ‘‘ Anisonyax? rufa’’ (p. 45).
RICHARDSON, J.
1829a. On Aplodontia, a new genus of the order Rodentia, constituted for
the-reception of the sewellel, a burrowing animal which inhabits
the northwestern coast of America. Zool. Jour., 4, 333-337.
Original description of genus, and of species leporina.
1829b. Fauna Boreali-Americana. (London, Murray), Part I, xlvi + 300
pp., 28 pls.
Descriptive remarks; genus Aplodontia (p. 210); species leporina
(pp. 211-213); cranium, foot, and tooth figured (figs. 7-14, pl. 18C,
opp. p- 197).
1837. Report on North American zoology, in Rep. 6th meeting Brit. Assoc.
Adv. Sei., 5, 121-224.
Apluodontia leporina listed (p. 150).
ScuHinz, H.
1845. Synopsis Mammalium (Solothurn, Jent), 2, 1-574.
Compiled account of Aplodontia leporina (pp. 138-139).
1918] Taylor: Revision of the Rodent Genus Aplodontia 493
SCHLOSSER, M.
1884. Die Nager des europiischen Tertiiirs. Palaeontographiea, 31, 19-161,
16 numbered, several unnumbered, figs. in text, pls. 5-12.
Abstract of Winge’s Jordfundne og nulevende Gnavere
(Rodentia) fra Lagoa Santa, Minas Geraes, Brasilien, E. Museo
Lundii, in Verzeichniss der anthropologischen Literatur, Archiv. f.
Anthrop., 19, 89-90.
Haplodontidae and Haplodon mentioned (p. 89); Haplodon (p.
90).
Scorr, W. B.
1913. A history of land mammals in the western hemisphere (New York,
Maemillan), xiv + 693 pp., frontisp., 304 figs. in text.
Aplodontia rufa mentioned (pp. 153, 233); Aplodontiidae (p.
249).
Srncuair, W. J.
1903. <A preliminary account of the exploration of the Potter Creek Cave,
Shasta County, California. Science, n.s., 17, 708-712.
“*Aplodontia either a new species or a new subspecies of 4.
major’’ listed (p. 711).
1905. New mammalia from the quaternary caves of California. Univ. Calif.
Publ. Bull. Dept. Geol., 4, 145-161, pls. 19-23, 2 figs. in text.
Original description of Aplodontia major fossilis (p. 147).
STEPHENS, F.
1906. California mammals. (San Diego, West Coast Publishing Co.), 351
pp., frontisp., many unnumbered figs. in text.
Family Aplodontidae, genus Aplodontia, and two species, major
and phaea, discussed (pp. 98-95).
Strong, W., and Cram, W. E.
1902. American animals. (New York, Doubleday), xxiii + 318 pp., many
unnumbered illustrations.
Family Aplodontidae and species Aplodontia rufa discussed (pp.
150-151).
SuCKLEY, G.
1860. Report, in Zool. Rep., U. S. Pac. R. R. Expl. and Surv., 12, bk. 2, pt.
3, no. 2, chap. 2, pp. 89-106.
Remarks on, and description and habits of, Aplodontia leporina
(p. 100).
SUCKLEY, G., and GrBBs, G.
1860. Op. cit., chap. 3, pp. 107-139, pls. 2, 5, 7, 9, 15.
Distribution of Aplodontia leporina; habits, Indian traditions
(pp. 124-126).
Swanson, W.
1835. On the natural history and classification of quadrupeds. (London,
Longmans), viii + 397 pp., 176 figs. in text.
Aplodontia (genus) discussed; Ap[lodontia]. leporina mentioned
(p. 388).
Taytor, W. P.
1914. A previously undescribed Aplodontia from the middle north coast of
California. Univ. Calif. Publ. Zool., 12, 297-300.
Original description of+ Aplodontia nigra.
494 University of California Publications in Zoology (Vou. 17
1916a. Aplodontia humboldtiana: a new mountain beaver from the Humboldt
Bay district, California. Proc. Biol. Soe. Wash., 29, 21-24.
Original description of Aplodontia humboldtiana.
1916b. The status of the beavers of western North America, with a con-
sideration of the factors in their speciation. Univ. Calif. Publ.
Zool., 12, 413-495, figs. A-Q in text.
Paragraph on the Aplodontiidae (pp. 470, 471). ‘
1916c. Two new aplodontias from western North America. Univ. Calif.
Publ. Zool., 12, 497-501.
Original deseriptions of Aplodontia rufa grisea (p. 497), and
Aplodontia californica columbiana (p. 499).
THOMAS, O.
1896. On the genera of rodents: an attempt to bring up to date the cur-
rent arrangement of the order. Proc. Zool. Soc. London, 1896,
1012-1028.
Places Anomalurus and Aplodontia in groups by themselves,
specifically providing that their true relationships will have to be
shown by further research (p. 1014).
TOWNSEND, C. H.
1887. Field-notes on the mammals, birds and reptiles of northern California.
Proc. U. 8. Nat. Mus., 10, 159-241.
Discussion of Haplodon rufus (pp. 174-175); mention made of
H. major (footnote, p. 174).
TROUESSART, FE. L.
1897. Catalogus mammalium tam viventium quam fossilium. (Berlin, Fried-
lander), tome 1, fase. 2, pp. 219-452.
Lists two species of Aplodontia, namely [ Aplodontia] rufa and
[Aplodontia] major (p. 450).
1904. Catalogus mammalium tam viventium quam fossilium. Quinquenalle
supplementum: Rodentia, fase. 2, pp. 289-546.
Lists seven forms of Aplodontia, namely [Aplodontia] rufa
[4. r.] californica, [A. r.] rainieri, [A.] pacifica, [A.] phaea, [A.]
olympica, | A.] major (p. 348).
TRUE, F. W.
1885. <A provisional list of the mammals of North and Central America,
and the West Indian Islands. Proc. U.S. Nat. Mus., 7, App., 587—
611.
Reference to family Haplodontidae and to Haplodon rufus
(p. 596).
TULLBERG, T.
1896. Zur Anatomie des Haplodon rufus. Zooliska Studier (Upsala, Alm-
quist), pp. 231-251, 2 pls.
Anatomy and remarks on classification (p. 250).
WAGLER, J.
1830. Natiirliches System der Amphibien mit vorangehender Classification
der Siugethiere und Vogel (Munich, Cotta), vi + 354 pp., 2 tables.
Haplodon listed (p. 4); sewellel (p. 22).
WaGner, J. A.
1843. Die Siiugethiere [Schreber]. Supplementband, 3. Abt., pp. xiv + 614.
Haplodon characterized; Haplodon leporinus discussed (pp. 395—
396).
1918] Taylor: Revision of the Rodent Genus Aplodontia 495
WALLACE, A. R.
1876. The geographical distribution of animals. 2 vols. (London, Mae-
millan), 2, viii + 607, 2 maps, 7 pls.
WEBER, M.
1904. Die Saugetiere (Fischer, Jena), xi + 866 pp., 567 figs. in text.
Aplodontia referred to ‘‘Familie’’ Haplodontoidea (p. 496).
WinGE, H.
1888. Jordfundne og nulevende Gnavere (Rodentia) fra Lagoa Santa, Minas
Geraes, Brasilien. E. Museo Lundii., 3, 1-200, pls. 1-8.
Haplodontidae (pp. 108, 110, 115, 135, 189); Haplodon (pp. 108,
115, 164, 189); relationships of rodent families (p. 110); Haplo-
dontidae thought to be derived from Ischyromyidae, and Anoma-
luridae and Sciuridae derived from Haplodontidae.
ZITTELL, K. A.
1894. Traité de paléontologie. I. Paleozoologie: 4. Vertebrata; mammalia.
(Paris, Doin), xi + 806 pp., 591 figs. in text.
Haplodontidae referred to Protrogomorpha (p. 523).
EXPLANATION OF PLATE 25
Crania of Aplodontia rufa phaea, to show some of the changes which take
place with age; X 0.67. Figs. la, 1b, 1c, no. 20317, Mus. Vert. Zool.; figs. 2a,
2b, 2c, no. 20306, Mus. Vert. Zool.; figs 3a, 3b, 3c, no. 8974, Mus. Vert. Zool.
Note in the crania dorsally the disproportionate expansion of the skull, par-
ticularly posteriorly; the marked reduction in width of the interorbital con-
striction; and the disappearance of sutures. In the crania ventrally note the
increase in measurements, the disappearance of sutures, and the disproportionate
lateral growth of the mastoid processes as compared with the audital tubes; but
note that the distance across the palate between the fourth premolars remains
nearly constant. In posterior view note the disproportionately rapid outgrowth
of the mastoid processes, with consequent disappearance of audital tubes, the
change in outline of the foramen magnum, and the change in outline of the
entire posterior aspect of the cranium, due to the development of the lambdoidal
ridges and the outgrowth of the mastoid processes.
[ 496 ]
pet
EXPLANATION OF PLATE 26
Dorsal view of crania of woodchuck, aplodontia, and beaver; all xX 0.50.
Fig. 4, Marmota flaviventris sierrae, no. 15165, Mus. Vert. Zool.; fig. 5, Aplodontia
rufa californica, no. 18663, Mus. Vert. Zool.; fig. 6, Castor subauratus subauratus,
no. 12654, Mus. Vert. Zool.
Note in Aplodontia and Castor the absence of postorbital processes on the
frontal; also the comparative width of the cranium posteriorly in Aplodontia,
as compared with Marmota and Castor.
[ 498 ]
ov
1%
|
.
|
|
|
|
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EXPLANATION OF PLATE 27
Figs. 7, 8.—Anterior view of crania of sciurid type and aplodontid type, to illus-
trate difference in infraorbital arrangement; X 0.77. Fig. 7, Marmota flaviventris
sierrae, no. 15165, Mus. Vert. Zool.; fig. 8, Aplodontia rufa californica, no. 18663,
Mus. Vert. Zool.
Note in Marmota the broadly expanded fossa for the masseter muscle just
laterad of the infraorbital foramen on either side; note also the compression and
reduction in size of the infraorbital foramen in this genus; note, on the other
hand, the generalized character of the region in Aplodontia, there being no
provision for the masseter muscle anterior of the ventral border of the anterior
root of the zygomatic arch, and the infraorbital foramen being comparatively
large, much as in the Eocene rodent family Ischyromyidae.
Figs. 9,10. Dorsal view of mandibles of aplodontid and sciurid, to illustrate
different development of angular processes; X 0.66. Fig. 9, Marmota flaviventris
sierrae, no. 15165, Mus. Vert. Zool.; fig. 10, Aplodontia rufa californica, no. 18663,
Mus. Vert. Zool. :
Note the lateral expansion of the angle in Aplodontia as compared with
Marmota.
[500 ]
EXPLANATION OF PLATE 28
Dorsal view of crania of three subspecies of Aplodontia; X 0.54. Fig. 11,
Aplodontia rufa rainieri, no. 90144, Biol. Sury. Coll.; fig. 12, Aplodontia rufa
columbiana, no. 1899, Mus. Comp. Zool.; fig. 18, Aplodontia rufa rufa, no. 3751,
Mus. Vert. Zool.
Note the general similarity in size between Aplodontia rufa rainieri and A. r.
columbiana, with the latter somewhat the larger; note the heavier zygomatic
arch in columbiana as compared with either of its neighbors, together with the
greater expansion of the arch near the posterior root. Note the tendency in
rainieri and columbiana to have temporal ridges closer together than in Aplo-
dontia rufa rufa. Note in the latter the general smaller size and lighter zygo-
matic arch as compared with the mountain subspecies.
[ 502 ]
LL
OE ———
EXPLANATION OF PLATE 29
Study of fresh specimen of Aplodontia rufa californica; male, no. 22618, Mus.
Vert. Zool.; East Fork Indian Cafion, Yosemite National Park, California;
June 19, 1915; X 0.35.
Note the small ear with white spot at base, the short tail, the chunky body
and blunt head.
[ 504 ]
Aqlvid [YOUAVL)
62
po
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UNIVERSITY OF CALIFORNIA PUBLICATIONS
IN
ZOOLOGY
Vol. 17, No. 17, pp. 505-515, 3 figures in text May 4, 1918
THE SUBSPECIES OF THE MOUNTAIN
CHICKADEE
BY
JOSEPH GRINNELL
(Contribution from the Museum of Vertebrate Zoology of the University of California)
Fieldwork was carried on by the California Museum of Vertebrate
Zoology during 1917 in the Inyo region of eastern California. In
going over the collection of birds obtained, the attention of the writer
was arrested by certain peculiarities evident in the Mountain Chicka-
dees. Comparison with series from the Sierras showed the Inyo birds
to be paler colored and longer tailed; and in order to appraise these
differences in taxonomic terms it became necessary to assemble ma-
terial representative of the entire range of the species, in so far as
possible. The results of the study thus undertaken are presented
herewith.
The material involved in the inquiry has amounted to 464 skins
of the Mountain Chickadee, derived from the following sources other
than the Museum of Vertebrate Zoology: United States National
Museum, through Dr. Charles W. Richmond; United States Biological
Survey, through Mr. Edward W. Nelson; and the private collections
of Messrs. Edward R. Warren, Joseph and John W. Mailliard,
G. Frean Morcom, Harry S. Swarth, and J. Grinnell.
As of general interest, and in the nature of an introduction to the
systematic analysis to follow, it may be stated that Penthestes gambeli
including its subspecies is throughout its range non-migratory, save
as a few individuals in pairs or small companies occasionally descend
in fall or early winter to lower levels closely adjacent to their moun-
tain habitats. The range of the species roughly extends from and
includes the Rocky Mountains to or nearly to the Pacifie Coast, and
from Alberta and British Columbia south nearly to the Mexican line—
somewhat south of it in northern Lower California. Within this gen-
eral area the Mountain Chickadee is by no means uniformly distrib-
506 University of California Publications in Zoology — [Vou. 17
uted. Especially towards the south is its range very ‘‘spotty,’’ the
representations on detached mountain tops being wholly isolated.
Two main areas of relatively continuous distribution are, however,
pereeivable—the Rocky Mountain area; and the Sierra Nevada area.
Close scrutiny of the series of specimens at hand well representing
the entire Rocky Mountain area reveals no variation in phylogenetic
characters from the northernmost to the southernmost stations. All
show in apparently equal degree the long tail and cinnamon tinge of
sides and back, these features together constituting the grounds for
separate subspecifie recognition of a Rocky Mountain form. On the
other hand, the Sierra Nevadan center, with its own recognizable race,
a~
Penthestes gambeli inyoensis |
Penthestes gambeli abbreviatus
Penthestes gambeli baileyae
Solid symbols, specimens examined
Hollow symbols, published records Bl
Breeding ranges outlined ]
DISTRIBUTION MAP
} MUSEUM OF VERTEBRATE ZOOLOGY
UNIVERSITY OP CALIFORNIA,
Fig. 1. Map showing distribution of the races of the Mountain Chickadee in
California.
1918] Grinnell: The Subspecies of the Mountain Chickadee 507
of relatively short tail, proves to have two outlying divergent forms.
These three forms are alike in their lack of any cinnamon tinge, this
being replaced in two of them by a buffy tinge and in one form by
leaden gray. The tail in one of the outlying forms is long, in the
other short. The habitats concerned are, respectively, the desert
mountains of the Inyo region of eastern California, and the coastal
mountains of southern California. This differentiation within the
Pacifie district, particularly within the state of California, will be
better understood in its geographic bearing by reference to the aecom-
panying map (fig. 1).
The behavior of the tail of Penthestes gambeli—long in the Rocky
Mountain district, short in the Pacific district (see figs. 2, 3)—is par-
alleled in the Penthestes atricapillus group of chickadees across the
North American continent in about the latitude of the state of Wash-
ee
—
Fig. 2. Tail of Penthestes gambeli gambeli (no. 27784, Mus. Vert. Zool.; ¢,
full-grown juv.; Sierra Ancha, Gila Co., Arizona; June 26, 1917). X 1.00.
Fig. 3. Tail of Penthestes gambeli abbreviatus (no. 24059, Mus. Vert. Zool.;
3, full-grown juv.; Yolla Bolly Mt., Tehama Co., California; August 5, 1913).
xX 1.00.
ington. In the northern Rocky Mountains oceurs the race P. a. sep-
tentrionalis, with long tail; in the Pacific Coast strip of Washington
and Oregon occurs the race P. a. occidentalis, with, among other char-
acters, relatively short tail. Other parallels are to be found in the
genera Psaltriparus, Thryomanes and Pipilo.
It is improbable that the fact of subspecific differentiation in
Penthestes gambeli has been altogether overlooked previous to the
present time. A premonition of the geographic variation now for-
mally pointed out is to be found in the early writings of Baird (1858,
p. 394), who remarked that ‘‘a specimen, apparently of this species
[Parus montanus|, from Medicine Bow ereek [Wyoming], (5643,)
though marked female, is larger than those from California, as shown
by the measurements. ’
in the subspecies now recognized are shown in the table of measure-
ments given by Ridgeway (1904, p. 409).
For a detailed description of the Mountain Chickadee as a species,
the reader is referred to Ridgway (1904, p. 408).
b)
Also, some of the differences in proportions
508 University of California Publications in Zoology | Vou. 17
Penthestes gambeli gambeli (Ridgway)
Rocky Mountain Chickadee
Parus montanus Gambel (1843, p. 259); and of authors, part. This name
preoccupied (see Ridgway, 1904, p. 409).
Parus gambeli Ridgway in American Ornithologists’ Union Committee
(1886, p. 335) ; and of authors, part. Substitute name.
Parus gambeli thayeri Birtwell (1901, p. 166). Based on adventitiously
colored specimen from Albuquerque, New Mexico. Type in United
States National Museum.
Penthestes gambeli, of authors, part.
Type locality—Santa Fé, New Mexico.
Diagnosis —Among the four subspecies of Penthestes gambeli here
recognized, color alone is sufficient for distinguishing P. gambeli gam-
beli. The flanks, sides of body and back in this form are pervaded
with a distinct tinge of cinnamon—more exactly, the ‘‘pinkish buff’’
of Ridgway (1912, pl. 29). In addition, this race shows the greatest
leneth of tail, and slenderest bill.
Measurements—Average of 14 males: Wing, 70.5 mm.; tail, 70.2;
exposed culmen, 8.6; depth of bill at base, 3.8. For extremes, see
accompanying table.
Note.—In this paper the length of wing is the usual chord of the folded wing
as dried at the side of the body in the conventional study skin. The tail length,
however, is measured from the base of the uropygium to the tip of the longest
pair of rectrices. It is found that in preparing specimens there is a practically
uniform place for cutting off from the body the uropygium with attached
feathers, and this is exactly where the tail bends back at an angle to the body in
the fresh bird. The lateral rectrices are rooted evenly with the base of the dried
uropygium so that the measurement of tail length thus taken becomes the distance
from the extreme proximal ends of the most lateral pair of rectrices to the tips
of the central pair—somewhat greater than the length of tail as given by Ridgway
(1901, p. xv), the latter being the measurement from the base of the central pair
of tail feathers to the tips of same. My reason for adopting this different pro-
cedure here is that more accuracy seems possible thereby, at least in the kind of
material here dealt with. There is less mussing of the specimen also.
Neither depth of bill nor culmen is a practical index to degree of
slenderness of bill. No ordinary method of measurement will suffice
to indicate the facts as they are perceived by the eye.
As will be seen by the dates in the tables, as a rule only unworn
specimens have been selected for measurement.
Range-—The Rocky Mountain region of North America, from
eastern British Columbia and western Alberta south to western Texas,
New Mexico and Arizona. Specimens examined, 72, from the follow-
ing localities:
Alberta: Smoky Valley, 50 miles north of Jasper House, 1; Henry House, 1;
15 miles west of Henry House, 1.
1918 Grinnell: The Subspecies of the Mountain Chickadee 509
British Columbia: South fork of Moose River, 1.
Montana: Gallatin County: Jefferson River, 1; Madison River, 2; Hillsdale, 2;
Mystic Lake, 1; Dry Creek, 1.
Idaho: Sawtooth Lake, 1.
Wyoming: Mammoth Hot Springs, 3; Jackey’s Creek, 4 miles southwest of
Dubois, 1; Teton Pass, 7200 ft., 2; Salt River Mts., head of Dry Creek, 9200 ft., 1;
Medicine Bow Mts., 10,200 ft., 1; Medicine Bow Creek, 1.
Utah: Filmore, 1.
Colorado: Loveland, 1; Middle Park, 1; Sangre de Christo Pass, 1; Fort Gar-
land, 1; Platte Canyon, 1; Pueblo, 2; Estes Park, 1; Boulder, 2; Gold Hill, 1;
Golden, 7300 ft., 1; Colorado Springs, 5; Querida, Custer County, 1; Salida,
Chaffee County, 1; Crested Butte, Gunnison County, 1; Stamford, 1.
Texas: Guadalupe Mts., 6800 ft., 1; Davis Mts., 1.
New Mexico: Fort Massachusetts, 1; Fort Wingate, 1; Albuquerque, 1; Taos
Mts., 8800 ft., 1; Manzano Mts., 4; Magdalena Mts., 7000 ft., 1; Cienequilla, 1;
Mt. Capitan, 1; Pecos Baldy, 1; Bear Spring Mts., 1; Shiprock, 1; Corona, 1;
Twining, 12,500 ft., 1; Fulton, 1; San Mateo Mts., 9500 ft., 1.
Arizona: Fort Whipple, 1; Mt. Graham, 1; San Francisco Mt., 2; Bright
Angel Spring, Kaibab Plateau, 1; Canyon Spring, 1; Santa Catalina Mts., 2;
Sierra Ancha, 6500 ft., 1.
Penthestes gambeli inyoensis, new subspecies
Inyo Mountain Chickadee
Parus gambeli, of authors, part.
Penthestes gambeli, of authors, part.
Penthestes gambeli baileyae, American Ornithologists’ Union Committee
(1910, p. 351), part (?).
Type locality—Panamint Mountains (northern part), 3 miles
east of Jackass Spring, 6200 feet altitude, Inyo County, California.
Type, male adult, no. 28782, Mus. Vert. Zool.; October 7, 1917; col-
lected by J. Grinnell, orig. no. 4588.
Diagnosis —The palest colored race of the four; sides, flanks and
‘cartridge
back, in unworn plumage, pervaded with pale buff—the
buff’? of Ridgway (1912, pl. 30). Wear or fading, or both, removes
most of this buff tone, so that the resulting effect, in spring and sum-
mer birds, is of an ashy tone of coloration, distinctly lighter than in
any of the other three subspecies, in same stage. It seems probable
that there is a paler tone to the underlying plumage parts and that
this becomes revealed by loss of the superficial pigment-bearing por-
tions through the gradual progress of feather abrasion. Inyoensis
shows nearly as long a tail as does gambeli. Its bill is somewhat
smaller.
510 University of California Publications in Zoology [Vou. 17
Measurements.—Average of 10 males: Wing, 71.1 mm.; tail, 69.3;
exposed culmen, 8.1; depth of bill at base, 3.7. For extremes, see
accompanying table.
Range.—The higher mountains of eastern California lying east
and southeast of Owens Valley, from the vicinity of the Mono Craters
and the White Mountains, in Mono County, south to the Panamint
Mountains, in Inyo County. Specimens examined, 50, from the fol-
lowing loealities, all in California:
Mono County: Mono Mills, 1; near Benton, 2; near Big Prospector Meadow,
10,000-10,500 ft., White Mts., 15; Cottonwood Creek, 9200 ft., White Mts., 1.
Inyo County: Silver Canyon, 7000-8000 ft., White Mts., 8; Roberts Ranch, 8300
ft., White Mts., 1; head of Black Canyon, 8000 ft., White Mts., 2; Independence,
3900 ft., 1 (vagrant); Mazourka Canyon, 8000-10,000 ft., Inyo Mts., 3; near
Jackass Spring, 6000-6200 ft., Panamint Mts., 14; Hanaupah Canyon, 7500-9000
ft., Panamint Mts., 2.
Penthestes gambeli abbreviatus, new subspecies
Short-tailed Mountain Chickadee
Parus montanus, of authors, part.
Parus gambeli, of authors, part.
Penthestes gambeli, of authors, part.
Type locality—Horse Creek, Siskiyou Mountains (near Seiad
Valley P. O.), Siskiyou County, California. Type, male, no. 119
(orig. no., in coll. J. Grinnell) ; December 12, 1901; collected by Mal-
colm P. Anderson.
Diagnosis —Tone of color on sides, flanks and back the same as in
inyoensis, though not quite so pale, namely, in fresh plumage, cartridge
buff. Tail (see figs. 2,3) much shorter than in either gambeli or inyo-
ensis ; and bill averaging smaller than in any of the other three races.
Measurements—Average of 14 males: Wing, 69.7 mm.; tail, 65.0;
exposed culmen, 7.9; depth of bill at base, 3.8. For extremes, see
accompanying table.
Range—The higher mountains of central and northern California,
southern Oregon (probably this subspecies), and northwestern Ne-
vada. Occurs west in northern California through the Siskiyou and
Salmon mountains and to South Yolla Bolly Mountain and Mount
Sanhedrin; and south in the Sierra Nevada to the vicinity of Mount
Whitney. Specimens examined, 182, from the following localities :
1918] Grinnell: The Subspecies of the Mountain Chickadee 511
California: Modoe County: Sugar Hill, 4; Warner Mts., 27. Siskiyou County:
Mt. Shasta, 3; Jackson Lake, 5900 ft., 5; South Fork Salmon River, 5000 ft., 1;
head of Rush Creek, 6400 ft., 2; Kangaroo Creek, 3; Castle Lake, 5400 ft., 1; Horse
Creek, Siskiyou Mts., 11; Beswick, 1. Trinity County: head of Bear Creek, 6400
ft., 4; head of Grizzly Creek, 6000 ft., 6. Tehama County: near South Yolla Bolly
Mt., 4. Mendocino County: near Castle Peak, 1; near Sanhedrin Mt., 4. Lassen
County: Eagle Lake, 6. Plumas County: Meadow Valley, 1. Nevada County:
Independence Lake, 3. Placer County: Summit, 1; Cisco, 6000 ft., 18; Blue Can-
yon, 4700-5000 ft., 12; Dutch Flat, 1. Eldorado County: Tahoe Valley, 2; Kyburz
Station, 1. Mariposa County: vicinity of Yosemite Valley, 13. Mono County:
Warren Fork of Leevining Creek, 9200 ft., 1; Williams Butte, 7500 ft., 1. Fresno
County: Bullfrog Lake, 10,600 ft., 6. Inyo County (not typical): near Kearsarge
Pass at 8500 ft., 1; Little Onion Valley, 7500 ft., near Kearsarge Pass, 1; Cotton-
wood Lakes, 11,000 ft., 7. Tulare County (not typical): Whitney Creek, 11,000
"ft., 3; Whitney Meadows, 9800 ft., 14; Olancha Peak, 10,000 ft., 1.
Nevada: Pine Forest Mts., Humboldt County, 13.
Penthestes gambeli baileyae (Grinnell)
Bailey Mountain Chickadee
Parus montanus, of authors, part.
Parus gambeli, of authors, part.
Parus gambeli baileyae Grinnell (1908, p. 29).
Penthestes gambeli baileyae, American Ornithologists’ Union Committee
(1910, p. 351), part (?); and of authors,
Type locality—Mount Wilson, 5500 feet altitude, San Gabriel
Mountains, Los Angeles County, California. Type in coll. J. Grinnell.
Diagnosis —Tone of coloration on sides, flanks and back distinctly
““smoke gray’’ of
plumbeous—more exactly, on sides and flanks the
Ridgway (1912, pl. 46), and on back near the
same authority (pl. 51). The tail in this race is short as in abbrevi-
ce
mouse gray’’ of the
atus, but the bill is long and heavy, averaging thicker through than
in any of the other three races.
Measurements.—Average of 14 males: Wing, 70.0 mm.; tail, 65.4;
exposed culmen, 8.5; depth of bill at base, 4.1. For extremes, see
accompanying table.
Range.—Higher mountains of southern California, from the
extreme southern Sierra Nevada in Tulare County, and the Santa
Lucia Mountains in Monterey County, south to the Cuyamaca Moun-
tains, San Diego County; also (probably this subspecies) in the San
Pedro Martir Mountains, in northern Lower California. Specimens
examined, 160, from the following localities, all in California:
512 University of California Publications in Zoology | Vou. 17
Monterey County: Headwaters of Big Creek, 2. Ventura County: Mt. Pinos,
2. Kern County: Kiavah Mt., 7000 ft., near Walker Pass, 2. Tulare County (not
typical): Taylor Meadow, 7000 ft., 8; Cannell Meadow, 7500 ft., 1; Pine Flat,
7500 ft., 1; Long Meadow, 7700 ft., 1; Sirretta Meadows, 9000 ft., 3; Trout
Creek, 6000 ft., 10; near Trout Creek, 7500 ft., 2; Jackass Meadow, 7750 ft., 3;
Troy Meadows, 8000 ft., 1. Los Angeles County: Mt. Wilson, 25; Buckhorn
Canyon, near Mt. Waterman, 2; near Pasadena, 9 (vagrants); Verdugo, 1
(vagrant). San Bernardino County: near Cucamonga, 1; San Bernardino Mts.,
35; Victorville, 3 (vagrants). Riverside County: San Jacinto Mts., 29; Thomas
Mt., 1; Santa Rosa Mts., 10. Orange County: Santa Ana Mts., 3000 ft., 1
(vagrant?). San Diego County: Julian, 1; Cuyamaca Mts., 6.
General remarks.—In making use of the intrinsic color characters,
which are important here in distinguishing subspecies, it is, of course,
essential that the effects of extraneous factors be taken into account.
Prolonged wear and fading evidently serve to weaken the intensity of
the color tones, more especially the buffy ones. Then, too, chickadees
seem peculiarly susceptible to discoloration by smoke, soot and charred
wood; for example, our series from Cisco and Blue Cain, stations
along the Central Pacific Railway over the Sierra Nevada, even though
taken in September and October almost immediately after completion
of the fall molt, are obviously more or less begrimed with soot. On
the other hand, the autumn- and winter-taken series from the Yosemite
region and from the Siskiyou Mountains are clean, and show their
intrinsic color tones to good advantage.
Intergradation undoubtedly connects the four races of the Moun-
tain Chickadee into a continuous series of forms. Abundant material
at hand from that portion of the Sierra Nevada immediately south of
Mount Whitney shows complete transition from Penthestes gambeli
baileyae to P. g. abbreviatus; in fact, many of the specimens ean only
be placed arbitrarily in one category or the other. Several examples
from the vicinity of Mono Lake, in Mono County, California, and
from along the west flank of the Sierras in Inyo County, insensibly
bridge the interval between P. g. abbreviatus and P. g. inyoensis, espe-
cially when considered in connection with the individual variation to
which each race is subject in about normal degree.
Material at hand from different parts of the Great Basin is unsat-
isfactory either in that it is seanty or because of the worn state of the
plumage. A summer-taken series of 13 Mountain Chickadees (nos.
8952-8964, Mus. Vert. Zool.) from the Pine Forest Mountains, Hum-
holdt County, Nevada, shows in color no approach to P. gambeli gam-
beli. In this respect it is like P. g. inyoensis, but the tail averages
nearly as short as in P. g. abbreviatus. Taking all features into
a
1918 ] Grinnell: The Subspecies of the Mountain Chickadee 513
account it seems best placed under abbreviatus. Fresh-plumaged fall
specimens from this locality would make determination more certain.
A specimen (jj, no. 547, Mus. Vert. Zool.) from Anthony, Baker
County, Oregon, taken October 16, 1907, might be referred to P. g.
abbreviatus; but the tone of color of back and sides is much darker
than usual in that form. It is certainly much darker than in P. g.
inyoensis. Its tail is but 65.3 mm. long. A skin (9, no. 18, Morcom
coll.) from Camp Harney, Harney County, Oregon, February 17,
1875, has a tail length of 67.7 mm., and in depth of color is about
intermediate between abbreviatus and P. g. gambeli. A specimen
(2, no. 186639, U. 8. Nat. Mus., Biol. Surv. coll.) from Fort Spokane,
Lineoln County, Washington, September 28, 1890, is almost identical
with gambeli in coloration; but it, too, has a short tail, only 64.4 mm.
long, though there is chance of error here, as several of the rectrices
are missing. The writer prefers to leave these last three examples for
the time being unplaced.
It is clear that intergradation between Penthestes gambeli gambeli
and any one of the other three subspecies is less well established than
between any two of these other three. But sufficiency of material from
the proper localities through eastern Oregon and eastern Washington
would likely prove its existence as is the case between the California
forms.
MEASUREMENTS (IN MILLIMETERS) OF SELECTED SPECIMENS REPRESENTING THE
Four Races or Penthestes gambeli
Penthestes gambeli gambeli
N. a Exposed Depth
no. Sex Date Locality Wing Tail culmen of bill
160697 ¢ Noy. 24,1892 Boulder, Colo. 71.7 71.2 8.8 4.0
109948 @ Oct. 8,1886 Pueblo, Colo. 72.0 72.4 8.4 3.9
109949 ¢ Oct. 5,1886 Pueblo, Colo. 70.9 70.3 8.5 4.0
176650 og Sept. 23,1888 Madison R., Gallatin Co., Mont. 68.0 67.8 8.9 4.1
124151 g Sept.15,1888 Jefferson R., Gallatin Co., Mont. 69.7 67.7 10.0 3.6
188823 ¢ Oct. 15,1902 Mammoth Hot Springs, Wyo. 68.1 67.0 9.0 3.6
62546 og Nov. 17,1872 Filmore, Utah 69.8 68.3 8.3 3.5
1366381 4 Sept.27,1890 Sawtooth Lake, Idaho 68.1 69.7 8.8 3.8
2282271 g Sept.14,1910 Teton Pass, Wyo. 71.2 70.4 8.3 3.6
1930861 ¢ Nov. 6,1903 Manzano Mts., N. Mex. 71.4 70.6 7.8 4.0
184653 ¢ Oct. 22,1902 Corona, N. Mex. 69.0 71.0 9.0 4.1
192942 g Oct. 10,1903 Twining, N. Mex. 72.0 71.4 8.5 4.0
1366371 g Sept.18,1889 San Francisco Mt., Ariz. 71.3 70.3 7.8 3.8
205661 ¢ Sept.10,1909 Kaibab Plateau, Ariz. 73.5 74.3 8.3 4.0
1 Biol. Sury. Coll.
514
Mus.
Vert.
Zool.
no.
28751
28760
28766
28767
28770
28771
28773
28774
28781
28782
+ Type.
QUWAKAE
OQ, Oy AY AY
=
No.
114
117*
119%
129%
164°
178*
244"
23307
23302
23298
25263
25800
26117
26342
n
is)
lal
&
Oy Oy Ay Oy Oy AY Oy Oy AY OG AQ AY OY OY
University of California Publications in Zoology
Date
July 24,1917
July 29, 1917
July 31,1917
Aug. 18, 1917
Sept. 29, 1917
Oct. 251917
Oct. 25,1917
Oct. 2,1917
Oct. 5,1917
Oct. 7, 1917
Date
9,1901
2. 12,1901
. 12,1901
. 14,1901
. 29, 1901
4, 1902
. 16, 1902
. 21,1912
- 19) Lo12
. 14,1912
. 26,1914
June 11,1915
Oct. 22,1915
Nov. 27,1915
Penthestes gambeli inyoensis
Locality Wing
White Mts., Mono Co., Calif. 73.0
White Mts., Mono Co., Calif. 71.0
White Mts., Mono Co., Calif. 72.4
White Mts., Inyo Co., Calif. 69.4
Panamint Mts., Inyo Co., Calif. 68.8
Panamint Mts., Inyo Co., Calif. 71.4
Panamint Mts., Inyo Co., Calif. 72.7
Panamint Mts., Inyo Co., Calif. 69.0
Panamint Mts., Inyo Co., Calif. 71.0
Panamint Mts., Inyo Co., Calif. 72.0
Penthestes gambeli abbreviatus
Locality Wing
Horse Cr., Siskiyou Mts., Calif. 68.4
Horse Cr., Siskiyou Mts., Calif. 69.0
Horse Cr., Siskiyou Mts., Calif. 70.0
Horse Cr., Siskiyou Mts., Calif. 68.4
Horse Cr., Siskiyou Mts., Calif. 69.0
Horse Cr., Siskiyou Mts., Calif. 66.8
Horse Cr., Siskiyou Mts., Calif. 70.8
Blue Cafion, Placer Co., Calif. 71.8
Blue Canon, Placer Co., Calif. 71.2
Blue Caiion, Placer Co., Calif. 71.6
Yosemite Park, Calif. 69.8
Yosemite Park, Calif. 72.8
Yosemite Park, Calif. 69.0
Yosemite Park, Calif. 67.1
[ Vou. 17
Exposed Depth
1 Collector’s original number, in Grinnell coll.; rest of numbers, Mus. Vert. Zool.
2\"T'ype.
No.
182?
7251
963+
6367
637?
1832?
18347
1836?
5516?
6073?
7458?
9958*
3364°
Xooco-
n
ro)
ss
Oy Oy A Oy Oy AY OY AQ AY OY QA AY OY AY
* Coll. H.
Date
Nov. 29, 1895
Nov. 1, 1897
Oct. 31,1898
Feb. 1, 1896
Feb. 1, 1896
. 12, 1896
. 12,1896
. 12,1896
Nov. 27, 1903
Sept. 21, 1904
Dee. 23,1905
Oct. 31,1897
Sept. 18, 1908
July 4, 1905
S. Swarth.
? Coll. J. Grinnell.
’Mus. Vert. Zool.
*Coll. J. & J. W. Mailliard.
5 Type.
Penthestes gambeli baileyae
Locality Wing
Mt. Wilson, Los Angeles Co., Calif. 67.0
Mt. Wilson, Los Angeles Co., Calif. 70.1
Mt. Wilson, Los Angeles Co., Calif. 67.0
Mt. Wilson, Los Angeles Co., Calif. 70.3
Mt. Wilson, Los Angeles Co., Calif. 68.6
Mt. Wilson, Los Angeles Co., Calif. 72.3
Mt. Wilson, Los Angeles Co., Calif. 65.4
Mt. Wilson, Los Angeles Co., Calif. 67.7
Mt. Wilson, Los Angeles Co., Calif. 72.1
Pasadena, Los Angeles Co., Calif. 69.7
Pasadena, Los Angeles Co., Calif. 70.0
Mt. Wilson, Los Angeles Co., Calif. 70.3
Santa Ana Mts., Orange Co., Calif. 70.8
Had. of Big Cr., Monterey Co., Calif. 70.5
Tail culmen of bill
72.4 29) 3.7
67.0 8.0 3.6
67.5 8.8 3.0
68.1 8.0 3.7
69.8 8.1 3.7
70.2 7.4 4.0
71.0 8.4 3.8
66.0 8.1 3.7
70.0 8.3 3.7
71.5 8.4 3.5
Exposed Depth
Tail culmen of bill
63.8 7.3 3.9
64.7 veo 3.7
65.0 7.8 3.6
63.0 8.0 3.6
64.2 7.2 3.9
63.0 7.2 3.7
65.0 7.3 3.6
69.5 8.2 3.9
64.3 8.8 4.1
65.7 7.8 3.5
65.9 8.4 3.8
67.7 8.7 4.0
65.0 8.8 4.0
63.6 8.3 3.9
Exposed Depth
Tail culmen of bill
63.0 8.9 4.3
66.9 9.0 4.4
63.3 8.2 4.0
65.5 8.3 4.1
65.8 8.2 4.0
65.7 fete 4.0
63.7 8.0 3.9
63.0 a 4.0
66.6 8.6 3.9
66.0 9.0 4.3
66.7 7.8 3.6
67.2 8.6 4.3
66.0 9.5 4.0
66.0 9.0 4.2
1918 | Grinnell: The Subspecies of the Mountain Chickadee 515
LITERATURE CITED
AMERICAN ORNITHOLOGISTS’ UNION COMMITTEE.
1886. The code of nomenclature and check-list of North American birds
adopted by the American Ornithologists’ Union. (New York, Amer-
ican Ornithologists’ Union), viii + 392 pp.
1910. Check-list of North American birds. Ed. 3, revised (New York, Amer-
ican Ornithologists’ Union), 430 pp., 2 maps.
Bairp, 8. F. (‘with the co-operation of’’ Cassin, J., and LAWRENCE, G. N.)
1858. Birds. Pacific Railroad Reports, 9, lvi + 1005 pp.
BirtWELL, F. J.
1901. Description of a supposed new subspecies of Parus from New Mexico.
Auk, 18, pp. 165-167.
GAMBEL, W.
1843. Descriptions of some new and rare birds of the Rocky Mountains and
California. Proc. Acad. Nat. Sci. Phila., 1, pp. 258-262.
GRINNELL, J.
1908. The southern California chickadee. Condor, 10, pp. 29-30.
Ripeway, R.
1901. The birds of North and Middle America. U.S. Nation. Mus. Bull., 50,
part I, xxx + 715 pp., 20 pls.
1904, Jbid., part III, xx + 801 pp., 19 pls.
1912. Color standards and color nomenclature. (Washington, D. C., Ridg-°
way), ii + 44 pp., 53 pls.
Transmitted March 3, 1918.
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“EXCAVATIONS OF BURROWS OFTHE RODENT
Di
HH
ree
APLODONTIA, WITH OBSERVATIONS ON
CHARLES LEWIS CA
MP
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IN
ZOOLOGY
Vol. 17, No. 18, pp. 517-536, 6 figures in text June 22, 1918
EXCAVATIONS OF BURROWS OF THE RODENT
APLODONTIA, WITH OBSERVATIONS ON
THE HABITS OF THE ANIMAL
BY
CHARLES LEWIS CAMP
(Contribution from the Museum of Vertebrate Zoology of the University of California)
CONTENTS
PAGE
TETRA NEYO A a ae eS ee A Ee ee 517
Historical
VIG) OP IE) 9 sneer Eek pee a
Burrows ..
LBD oese See et aeons eee gt RSE Oa eT eee ees
TRS bib vey LOE EY aoe ce ee ee Sac a BRP eee 528
Hibernation
Sc atol oprymsevmuntert1 OVI p aN A Care BNy BO Oye eae m eas soa aa tee ence ane eee cena ena 531
PSI STINS (2) OSLO OR OTE cae eer cr Oo RO occa eee 532
Breeding habits ....
AVC tec ce eae et ss Dig Lac setae nae econ wn neces Noe vunan pictse beta dbdecdecediaence eee tees 538%
NB TT @ VENT GS apres aoe an cts «2s oc ce reece tevez az we ccucaecanevetavet oe sve aes egsbe tes aera ateneere Bee ote 533
Economie importance and trapping ..
MebeTATUT OM CUGG Cees stern nee sneer cen cree en:
INTRODUCTION
While engaged in field work for the University of California
Museum of Vertebrate Zoology the writer has had the opportunity
of observing personally in the wild three of the California races of
Aplodontia or Mountain Beaver. The notes of other members of
Museum field parties have been available also in the preparation of
this report, in fact all material, from whatever source, in the files
of the Museum of Vertebrate Zoology. Im addition, Mr. Alfred C.
518 University of Califorma Publications in Zoology | Vou. 17
Shelton, of the University of Oregon, furnished information which
has been made the basis of some of the statements herein contained.
The writer wishes to thank Drs. Joseph Grinnell and Walter P.
Taylor for bringing this paper into final form for publication.
Historica
Present knowledge of the mountain beaver began with the arrival
upon the coast of Oregon of the expedition under Lewis and Clark.
This expedition went into winter quarters at Fort Clatsop, near the
present site of Astoria, on December 8, 1805, In Clark’s notes (in
Lewis and Clark, 1904, 3, p. 279) of Friday, December 13, 1805,
appears the following entry:
‘*The Clatsops leave us to day after a brackfast on Elk which they
appeared to be very fond of before they left us they Sold me two robes
of the skins of a Small animal about the size of a cat, and to Captain
Lewis 2 Cat or Loucirva Skins for the purpose of makeing a Coat.’’
The ‘‘small animal’’ mentioned was what the explorers thought the
Indians ealled ‘‘Sewelel,’’ if we are to judge by the notes of Lewis
for February 15, 1806 (loc. cit., 4, p. 73), and February 26, 1806
(loc. cit., pp. 109-110).
The first published account of the mountain beaver was a modifica-
tion of the above notes in the Biddle-Allen edition (1814) of the Lewis
and Clark Expedition (see Coues, 1877, p. 591). And the first nat-
uralist to study a skin of the animal was Richardson (1829, pp.
211-213).
The Wilkes Exploring Expedition collected two specimens of
aplodontia, probably in the neighborhood of Puget Sound. T. R.
Peale (see Coues, loc. cit.), who collected these specimens and later
(1848) described them, remarks on the abundance of the species at
““Puget’s Sound.”’
The next notice of the genus was by Audubon and Bachman
(1854, 3, pp. 99-102), who quote Richardson and Lewis and Clark as
first published, and add inferences drawn from the anatomy as to the
habits, and some further statements concerning distribution.
The naturalists of the Pacifie Railroad Survey, under the super-
vision of 8S. F. Baird, were able to make valuable observations on the
habits of aplodontia. Their findings together with those of other
observers since then and up to the year 1916, are incorporated into
the sueceeding parts of the present paper.
1918 | Camp: Burrows of the Rodent Aplodontia 519
Coues (1877, p. 591) reviews the literature on Aplodontia up to
his time, making, however, no mention of Lord’s interesting notes on
the genus (1866, 1, pp. 346-358).
The most important recent contribution to the life-history of aplo-
dontia is comprised in the notes and photographs of Anthony (1916).
This came to hand after the present paper had been prepared.
The writer has not had access to an article by Murphy (1876), of
which he has heard.
For full systematic treatment of the genus Aplodontia and for
lists of names, both vernacular and scientific, see Taylor (1918).
Hapirat
Like the beaver and some of the ground squirrels, the aplodontia
seems to be of social habits, congregating in more or less extensive
colonies in localities where conditions favor its existence. If a real
social bond exists it must be a primitive one. The scattered nature of
most of the colonies proclaim this; and the probable absence of voice,
which usually accompanies social habits, is suggestive.
The factors necessary for the existence of all the nine subspecies
of Aplodontia now recognized seem to be: An abundant supply of
certain food plants chiefly of riparian growth; and a soil at least
moderately deep and firm, and with adequate drainage, where the
animals can burrow. Hillside seepage areas and the banks of streams
provide the usual situations of colonies. Colonies occurring in wooded
and brushy places at a distance from water are sometimes inhabited
only in the wet season (Shelton, MS, 1916), but if the food supply
continues, such burrows may be used the year. round.
Burrows
Wherever the aplodontia lives it digs extensive underground tun-
nels that in a populous colony form a network of passages a few
inches beneath the surface of the ground (see fig. 1). Each burrow
system has many openings to the surface, but excavated dirt and
rubbish is pushed out usually at only a few of these holes. The three
subspecies living along the coast of California invariably choose for
the colony a spot within a dense tangle of ferns, thimbleberry and
other low plants in wet or at least damp soil. So well concealed
beneath tangled shrubbery are the unplugged burrows that the pres-
520 University of California Publications in Zoology | Vou. 17
ence of the animals is often unknown to persons resident in even the
immediate neighborhood.
The grouping of the burrows in close ‘‘companies’’ may simply
indicate that the animals are not wont to range widely enough to
‘
invade adjacent districts, though these seemingly might support a
population of similar extent. Overcrowded conditions may prevail in
one place, while territory of the same character remains unoccupied
near by. Near Point Reyes, in the gulches five miles west of Inver-
ness, Marin County, California, trapping was carried on in 1913 in
two colonies of Aplodontia rufa phaea. The burrows extended along
UNOPENED ~<TUNNEL
4-24
SCALE
——————
ONE METER
ooo
THREE FEET
Fig. 1. Plan of underground tunnels in colony of Aplodontia rufa phaea
excavated near Point Reyes, Marin County, California. All tunnels shown were
within 600 millimeters of surface of ground.
the side of a low north-facing bluff among bare hills for two miles or
more. One section of this colony in a rectangular area about 500 by
100 feet contained at least 100 burrow entrances. Here 169 ‘‘trap-
nights’’ (one trap out one night equals one trap-night) in thirty-six
burrow entrances produced eleven specimens. No individuals were
secured in the last forty-six trap-nights, an indication, perhaps, that
the catch approximated the total population of that part of the colony.
Ninety-two trap-nights in another limited part of the above colony
produced six specimens.
Some of the burrows in the Point Reyes locality were excavated by
me (see figs. 1, 2). The floors of many of the runways were seen to be
tramped hard from long use. It was found that nearly all the burrow
1918 ] Camp: Burrows of the Rodent Aplodontia 521
entrances on the hillside were connected with one another by passages
six to eighteen inches beneath the surface. At one point a nest was
discovered half fillmg a globular chamber about a foot in diameter
(see fig. 2). The nest was constructed for the most part of dry
sword fern ‘‘leaves’’ stripped from the frond stem. It also contained
a few large dry leaves of the cow parsnip, a plant nearly everywhere
found in the vicinity of the burrows. Some dirt was mixed with the nest
and the whole had been trampled down into a firm mass, flat on top.
EARTHEN PELLETS £
STORED Foo0 6
450-0
re
ONE METER
TAREC Feet
Fig. 2. Plan and partial elevation of nest-galleries of Aplodontia rufa phaea
excavated near Point Reyes, Marin County, California. Numbers on plan
indicate depth in millimeters beneath surface of ground.
Adjoining the nest chamber in the excavation mentioned, and opening
into it, was a low square ‘‘room’’ twenty inches in diameter, the floor
and sides of which showed signs of continued use by the inhabitants.
Pockets near by contained stored roots, stems and leaves carefully
blocked in by artificial earth pellets (see fig. 2).
Aplodontia rufa nigra, at Point Arena, Mendocino County, Cali-
fornia, the closest relative of A. r. phaea, and A. r. hwmboldtiana, of
the Humboldt Bay region and farther north, both apparently have
burrow systems similar to that of phaea described above. Like phaea,
mgra inhabits thimbleberry thickets. It lives on north-facing guleh
sides in a small area (about twenty-four square miles) along the cen-
522 Unwersity of California Publications in Zoology — (Vou. 17
tral Mendocino County coast. The colonies now known extend from
the town of Point Arena to Alder Creek, seven and one-half miles
north. In the case of the race phaea colonies oceur in a part of the
Point Reyes Peninsula and country adjacent, in Marin County, Cali-
fornia, an area of not more than 110 square miles in all.
Of a colony of Aplodontia rufa humboldtiana discovered at Cud-
deback, Humboldt County, California, Stephens (MS, 1910) says:
I put ten traps in the burrows and did not cover more than half the
entrances. I saw at least two tunnels run close under the surface, with fre-
quent openings. My previous idea of a burrowing place for aplodontia is wet
or springy land. This is far from that. It is a very steep south-facing slope
almost at the top of a point on a high ridge and is as dry a place as well
could be found. The location is in an open place in a thick forest of redwood,
spruce, and fir. It has been burned over three years ago and there is very
little underbrush.
Lyon (1907, p. 91), speaking of Aplodontia rufa rainieri, says:
““The burrows or tunnels are about 8 to 9 inches in diameter and were
always found in groups or colonies.’’
The habitat preferences and burrowing habits of Aplodontia rufa
rufa appear to be similar to those of the California coast species.
Matteson (1877, pp. 4384-435) states that the mountain beaver of
Oregon ‘‘is a digger par excellence and burrows into the sides of the
hills usually in the neighborhood of a spring.’’ Lum (1878, pp.
10-13) notes that the species of mountain beaver along the western
base of the Cascade Mountains ‘‘usually selects the open glades of the
forest, thickly grown up with fern and sallal’’ (Gaultheria shallon)
for its burrows. ‘‘Here the ground will be seen perforated with
holes. . . . Beneath the ground the various openings connect and form
a perfect plexus of passages often nearly parallel with the surface
and only a foot or so in depth. . . . In many places I found water
coursing its way through the passages.’’
In California, at the southern part of its range, aplodontia usually
prefers a shady, north slope for its burrows; but Brooks states (1899,
pp. 258-259) that the race (Aplodontia rufa columbiana) in British
Columbia, in the extreme northern end of the range of the genus,
prefers a locality ‘‘where there is a good thick growth of vegetation
on the mountain slopes, especially on those with a south exposure.’’
Aplodontia rufa californica of the Sierra Nevada, in California,
has been found in a variety of situations, at elevations all the way
from 5500 to 10,000 feet. Townsend (1887, pp. 174-175) saw burrows
near Morgans Springs near Mount Lassen, and on the North Fork of
1918] Camp: Burrows of the Rodent Aplodontia 523
the Feather River at Big Meadows. He says the burrows resembled
those of muskrats and were in clayey banks. On Mount Shasta,
Osgood (in Merriam, 1899, p. 93) found at most only a single pair
of mountain beavers living in one place.
Price (1894, p. 328) notes that at the big-tree grove near Red
Point, Placer County, ‘‘two small colonies were seen in bogey land
about springs.’’ Burrows were noted ‘‘in the bottom of a ravine
among dense beds of moss, thickly shaded with tangled bushes.’’
Stephens (1906, pp. 94-95) speaks of burrows of California
SCALE
—
ONE METER
7 —
PROFILE THROUGH LINE A-D *.” TnREE FEET
Fig. 3. Plan and partial elevation of burrow system of Aplodontia rufa
californica exeavated at Chinquapin, Yosemite National Park, California.
mountain beaver that were fifty yards or more in length. He observed
the animals on the headwaters of the Carson River in Alpine County.
Here, he says, they ‘‘live in wet springy land in canyons and on
mountain sides where suitable springs occur, usually at considerable
altitudes’’ [8000 feet]. Most of the entrances were under clumps of
willow. Stephens also saw where watercourses had been diverted by
the burrows.
Muir (1909, p. 201) deseribes how Aplodontia rufa californica
in the Yosemite National Park ‘‘digs canals and controls the flow of
small streams under the sod’’; ‘‘and it is startling,’’ he says, ‘‘ when
524 University of California Publications in Zoology [ Vou. 17
one is camped on the edge of a sloping meadow near the homes of
these industrious mountaineers to be awakened in the still night by
the sound of water rushing and gurgling under one’s head in a newly
formed eanal.’’
Grinnell (MS, 1912) observed Aplodontia rufa californica at Blue
Canon, Placer County, California. His notes in regard to their bur-
rowing habits follow. ‘‘It [the colony] is now [August 30] in rather
?
Fig. 4. Type of habitat in which ‘‘colony’
was situated; 9700 feet altitude, head of Lyell Caion, Yosemite National Park,
California. The trees are lodgepole pines; the riparian growth is willow.
of Aplodontia rufa californica
dry ground, though evidently wet most of, fhe year; for it is overgrown
with luxuriant thimbleberry and ee The whole place is
densely shaded by black oaks and spruces. At the colony located I
saw fully 15 fresh burrows.’’ There were evident trails or surface
runways out into the salmonberry bushes. Burrows were frequently
coincident with water channels beneath the sod.
In the Yosemite National Park extensive thickets of the preferred
food plants are scarce, and here the mountain beaver must as a rule
be content with limited tunnel systems through the narrow willow-
1918] Camp: Burrows of the Rodent Aplodontia 525
fringes along the streams. At Chinquapin, 6200 feet altitude, in the
Park, a tunnel (see fig. 3) of Aplodontia rufa californica was exca-
vated for a distance of twelve yards along the steep bank of a stream
without the discovery of a single side branch of importance. Only
two individuals, male and female, were captured in this tunnel. The
burrows in the colony at Chinquapin were far scattered along-stream
owing to the narrowness of the area available to the animals. At
many other localities in the Park similar conditions prevailed (see
fig. 4). Small colonies with more complicated systems of runways are
sometimes established where conditions will allow, as, for example, in
willow thickets bordering meadows and, high up in the mountains, in
Labrador tea and willows along creeks and about springs. In almost
every colony tunnels were found containing running water. At one
place the entire flow of a small stream was taken care of for some
distance below ground.
The burrows of Aplodontia rufa rufa of the Trinity and Siskiyou
mountains, to Judge from the descriptions given by Kelloge (1916,
pp. 369-372) and Alexander (MS, 1911), appear to be similar to the
more simple type of tunnel system of A. r. californica. Even in one
exceptional case where the ground was found to be ‘‘literally honey-
(a3
combed’’ by burrows and runways, a ‘‘main tunnel’’ could be dis-
tinguished. As with californica never more than one pair of animals
was secured from a single set of runways.
Foop
A wide range of plant species is eaten by the mountain beaver.
The number used for food at a single locality generally includes most
of the species of shrubs and young trees within reach of the burrows.
However, a marked preference is often shown for one particular
plant. Those kinds with succulent stems, when available, take prece-
dence over the more woody ones. The coast species of aplodontia are
predominantly fern and root eaters. The following table provides
record of the plants known to be eaten by various races of the moun-
tain beaver.
526 University of California Publications in Zoology
[ Vou. 17
TABLE SHOWING Foop of APLODONTIA ACCORDING TO SUBSPECIES AND LOCALITY
Authority
Lum (1878), p. 11)
Shelton (MS, 1916)
Alexander (MS,
1911)
Kellogg (1916, pp.
371-372)
Elliot (1899, p.
252)
Brooks (1899, pp.
258-259)
Heller (in Elliot,
1903, p. 184)
Anthony (1916,
p- 57)
Shelton (MS, 1916)
Stephens (MS,
1910)
Camp (MS, 1913)
Dixon (MS, 1909)
Camp (MS, 1913)
Camp (MS, 1913)
Merriam (1899b,
pp. 92-93)
Locality
Aplodontia rufa rufa
Oregon (?)
Vicinity of McKenzie Bridge,
Lane Co., Oregon
South Fork, Salmon River,
Siskiyou Co., California
Jackson Lake, Trinity Co.,
California
Aplodontia rufa olympica
Olympie Mountains, Wash-
ington
Plants
Ferns, salal, hazel, red
clover, and roots
Mountain alder, ‘‘wild pea’’
Cut grass (‘‘hay’’), and
green plants
Alder and wild plum
A kind of lily
Aplodontia rufa columbiana
Southern British Columbia
Aplodontia rufa pacifica
Gardiner and Florence, Ore-
gon
Tillamook, Oregon
Oregon (west-central)
Scotch cap (Rubus nutkanus)
Ferns, salal, and Oregon
grape
Thimbleberry, elk-brake, and
sword-fern
Salal, Oregon grape, vine
maple, young firs (up to
34 in. in diameter), sword-
fern (‘‘favorite food’’),
and wild rose
Aplodontia rufa hwmboldtiana
Cuddeback, Humboldt Co.,.
California
Aplodontia rufa nigra
Point Arena, Mendocino Co.,
California
Aplodontia rufa phaea
Lagunitas, Marin Co., Cali-
fornia
Five miles west of Inverness,
Marin Co., California
Three miles south of Olema,
Marin Co., California
Laurel sprouts, ferns, and
grass
Sword-fern, cow - parsnip,
stalks of nettle, cultivated
‘‘eow-peas and oat hay’’
Roots of mint
Sword-fern (fronds and root-
stocks), cow-parsnip, sal-
monberry, wild pea-vine,
and chilicothe (leaves and
roots)
Sword-fern and brake fern
Aplodontia rufa californica
Mt. Shasta, California
Ferns, willows, thimbleberry,
mountain ash, and brake
fern
1918 | Camp: Burrows of the Rodent Aplodontia 527
Authority Locality Plants
Townsend (1887, Feather River, California Weeds and coarse grasses
pp. 174-175)
Grinnell (MS, Blue Canyon, Placer Co., Choke-cherry, willow, creek
1912) California dogwood, and _ thimble-
Allen (in Merriam,
1886, pp. 312-
328)
Price (1894, pp.
315-332)
Stephens (1906, pp.
94-95)
Grinnell (MS,
1915)
Camp (MS, 1915)
Camp (MS, 1915)
Grinnell (MS,
1915)
Storer (MS, 1915)
Camp (MS, 1915)
Camp (MS, 1915)
Placer Co., California
Big-tree grove, 20 miles
southeast of Red Point,
Placer Co., California
Headwaters of Carson River,
8000 ft., Alpine Co., Cali-
fornia
West Fork of Indian Creek,
7250 ft., Yosemite Na-
tional Park, California
Head of Porcupine Creek,
above Poreupine Flat, 8400
ft., Yosemite National
Park, California
One mile northeast Porcu-
pine Flat, 9500 ft., Yo-
semite National Park, Cal-
ifornia
Porcupine Flat, 8600 ft.,
Yosemite National Park,
California
Head Lyell Canyon, 9800 ft.,
Yosemite National Park,
California
Chinquapin, 6200 ft., Yosem-
ite National Park, Cali-
fornia
One-half mile west Ostrander
Rocks, 7500 ft., Yosemite
National Park, California
berry (Rubus parviflorus)
Lily stems, willow, red osier,
small fir trees, and man-
zanita
Mountain cranberry stems
(Vaecinium — occidentale),
Ceanothus, and Rhododen-
dron, in large quantities
Iris, Astragalus, willow, and
alder
Oct. 30, large quantities of
“*hay’’ out to dry consist-
ing entirely of Lupinus
longipes
Willow and aspen (limbs up
to % inch in diameter)
Labrador tea (Ledum glandu-
losum), and Ribes visco-
sissimum
Willow bark, and young
lodge-pole pines (terminal
twigs only)
Willow stems up to % inch
in diameter
Azalea (stems to 1 inch in
diameter chewed), Ribes,
Cornus pubescens, chinqua-
pin, incense cedar, hazel,
Prunus (sp.?), Ceanothus
cordulatus, white fir, sugar
pine, brake fern
Cornus pubescens, and Ribes
viscosissimum.
In captivity the animal has been known to eat ‘‘apples and other
fruit and vegetables’’ (Lum, 1878, pp. 10-13), celery, carrots, turnips,
potatoes, cabbage, and lettuce.
528 University of California Publications in Zoology | Vou. 17
Frepine Hairs
Aplodontia is rarely seen in the daytime, but comes forth early in
the evening in search of food and doubtless remains active through-
out the night. Bretherton (1895, p. 124) mentions that he has trapped
a large number without ever having seen one abroad in daylight.
Our own observations as well as those of others confirm him. An indi-
vidual of Aplodontia rufa phaea that the author kept captive built
itself a nest where it remained passive in a sitting posture throughout
the day.
rs
4
ip
bf
a*/-
Fig. 5. Entranee to burrow of Aplodontia rufa phaea; six miles west of
Inverness, Marin County, California. Cutting of sword fern to be seen in mouth
of burrow.
This rodent is slow and cumbersome in gait and seldom ventures
far from its burrow. Tunnels are dug directly into suitable clumps
of vegetation where the animals wish to forage, and from the burrow
entrances issue short trails or runways along which the animal trans-
ports cut stems and leaves to the mouths of its burrow system. Occa-
sional sections of eut material are dropped along these pathways,
thus distinguishing them from the runways made by rabbits and
eround squirrels (see fig. 5). Remains of fresh leaves and twigs
1918] Camp: Burrows of the Rodent Aplodontia 529
pulled just within the burrow entrances lead one to believe that the
timid mountain beaver does much of its feeding in the comparative
security of its burrow.
A singular habit has been noticed in connection with the storage
of food. In a burrow excavated at Point Reyes (see fig. 2) the
entrances of two of the food storehouses were found plugged with
large pellets of earth evidently manufactured by the animal for this
especial purpose. These earthen balls were one to two inches in diam-
eter and very hard and dry, evidently from having been handled a
good deal. It is curious that the outer burrow entrances are not
similarly plugged.
Gibbs (in Suckley, 1860, p. 100) was the first to note a peculiar
hay-making habit among the mountain beavers he found living at the
top of Yakima Pass, Washington. ‘‘Near their abodes,’’ he says,
“were small bundles of some herb or plant eut with nicety and laid
out on logs to dry or wilt,’’ and Lyon (1907, p. 91) and Bretherton
(1895, p. 124) confirm this. Townsend (1887, pp. 174-175) thought
he recognized the hay-making instinet among the aplodontia he
observed on the North Fork of the Feather River in California.
Elliot (1899, p. 252) makes particular notice of the hay-making
habit and adds that the cured stems of the lily, which he saw Aplo-
dontia rufa olympica using, may be employed either for food or as
bedding in the nests. Stephens (1906, p. 95) writes: ‘‘I saw bunches
of plants laid up on low bushes to dry, commonly over entrances to
burrows, most of these not being much dried, as if they had carried
them in as soon as they were well wilted’’—this in regard to Aplo-
dontia rufa californica, on the headwaters of the Carson River in
Alpine County, California. Miss Kellogg (1916, p. 372) speaks of
the habit in question as occurring among the colonies of Aplodontia
rufa rufa discovered on the South Fork of the Salmon River in Siski-
you County, California. She says: ‘‘We found the mountain beavers
making what we called ‘hay’—large bunches of green plants of various
kinds cut up and spread out as if to dry and to be used later.’’
From observations of the nests and of feeding habits my own con-
clusion is that the animal does not eat dried food but uses its hay
for nest-building material.
The manner of feeding has been observed in the captive animal
and partly determined from the evidence of cuttings. The rodent
usually sits back upon its short tail, with the back curved, the hind
feet extending out in front, soles forward, and the fore paws grasping
530 University of California Publications in Zoology {Vou.17
the stem, root or leaf. The short first finger is used like a human
thumb and the large callosities on the palms aid in holding an object
so firmly that it can be retained easily in one hand. Stephens (1906,
p- 95) says that the hind feet as well as the fore feet are used in
erasping. Plants are harvested in sections from two to six inches in
length so that they can be dragged into the burrows. Tough woody
twigs and stems are chewed through from one side only, not in the
manner in which beavers attack large trees. If the plant is a soft
annual the whole stem is rapidly nibbled; if a perennial, the bark is
chewed off in the manner of a beaver or porcupine and the wood
discarded.
More than one writer since Lewis and Clark have mentioned the
climbing habit of mountain beavers; and the evidence seems to be
good that at least the Oregon and Washington coast species climb into
low bushes for food (see Lum, 1878, pp. 10-13). The mountain
beavers on the grounds of the University of Washington at Seattle
have been seen climbing.
HIBERNATION
There seems to be no certain evidence that the mountain beaver
hibernates even in the coldest part of its range. Cooper (1860, p. 82)
affirms that these animals had been seen running over the snow in
the Nisqually Valley, Washington. Suckley (in Suckley and Gibbs,
1860, p. 124) learned from the Nisqually Indians that the mountain
beaver moves about a little during the winter but does *‘
decidedly active till late in the spring.’’ Lord (1866, pp. 346-358)
maintains that the animal only partially hibernates—this of the
species in British Columbia. Matteson (1877, pp. 434-435) believes
that the mountain beaver hibernates, at that time covering the
entrance of the burrow with stalks of leaves and fern. Bretherton
(1895, p. 124) says that they do not hibernate.
Lum (1878, pp. 10-13) reports the following of the Oregon and
Washington species: ‘‘They do not hibernate, but keep their burrows
open all winter; beaten trails in the snow are often seen, leading
not become
above ground for a few feet from one hole to another. They are able
to gather their food at any time of the year, seldom going more than
a few feet from the entrance of their holes to procure it.”’ Brooks
(1899, p. 259) asserts that the aplodontia in British Columbia ‘‘does
not hibernate at lower levels but must do so on the higher mountains,
as it does not seem to make ‘hay’ like the Pika.’’
1918 | Camp: Burrows of the Rodent Aplodontia 531
The California species of mountain beaver, even in the high
mountains, are certainly active during the winter. Near Olema, in
Marin County, Aplodontia rufa phaea has been trapped, at its burrow
entrances, during the latter part of November. In the Yosemite
National Park, at elevations of from 6200 to 8600 feet, winter work
of Aplodontia californica has been observed—young pine, fir and
cedar trees with the limbs trimmed off up to ten feet above the
ground and with twigs trimmed from one side of the trunk only, as
if worked on when the trees were bent down in the snow. Azalea
Fig. 6. Life study of Aplodontia rufa californica; Lyell Canon, Yosemite
National Park, California; July 17, 1915. Note the small eye and ear, the white
spot at base of ear, the shortness of the tail, and the seemingly sluggish general
apearance of the animal. The matted appearance of the hair on the back is
due to moisture.
and other thickly growing bushes with the tops trimmed off evenly
a foot or more above the ground indicate that the animals probably
forage on top of the snow.
ScaroLoGy, SANITATION, AND DRAINAGE
The faeces are black, about the shape of those of a pocket gopher
but larger(in Aplodontia rufa phaea, measuring 6 to 7 by 15 to 20
millimeters) ; the urine is yellow and cloudy. Faeces are rarely seen
about the burrows; they are probably deposited in some particular
place underground, but I have never succeeded in discovering any in
the burrow systems excavated.
582 University of California Publications in Zoology — | Vou. 17
Arrangements are always made for the proper drainage of the bur-
rows, and in wet hillsides water often runs continuously from the
lower tunnels. In some places the courses of streams are diverted,
though perhaps accidentally. The mountain beaver is not aquatic in
any true sense, but is often caught in traps set in the water running
through the burrows. The animal is said to dip its fore feet in water
and to wash its head.
SENSE PERCEPTION
Sight and hearing seem to be relatively defective in the mountain
beaver (see fig. 6). The individual comes in contact with its under-
ground environment chiefly through the sense of feeling, and this is
markedly developed all over the body. The slightest touch upon a
hair will be responded to instantly by quick jerking movements. The
sense of smell seems to be good, for the animal has a way of frequently
raising its nose to sniff; also the scent glands are well developed.
BREEDING Hasits
Almost nothing is known concerning the breeding habits of any
of the forms of this interesting genus. Lord (1866, pp. 346-358),
speaking of Aplodontia rufa columbiana as observed upon the banks
of the ‘‘Chilukweyuk River’’ in British Columbia, asserts that the
female has from four to six young at a birth and about two litters a
year; and that the nest is like a rabbit’s, made of grass and leaves,
and is placed at the end of a deep burrow. Lum (1878, pp. 10-13)
writes that ‘‘ People living in the vicinity of these animals [A. r. rufa ?,
in Oregon ?] tell me that the young show’tls just weaned make their
appearance during the month of June in numbers from three to five
at a birth.”’
VOICE
The only sound that we have known the mountain beaver to make
is a singular grating noise produced when the animal is alarmed, by
rasping the lower incisors laterally across the tips of the upper. I
have seen pocket gophers, marmots, and copper-headed ground squir-
rels (Callospermophilus) do the same when terrified.
Aplodontia has been eredited with a real voice, but the chances
for error in most of the following reported observations in this regard
are very great. Suckley and Gibbs (1860, p. 124) quote a certain
Colonel Simmons as saying that he had seen mountain beavers ‘‘sitting
1918 | Camp: Burrows of the Rodent Aplodontia 533
at the entrances of their burrows early in the morning and whistling,
something in the manner of the prairie-dog.’’ Lord (1866, pp. 346—
358) heard whistlings in the evening which he attributed to these
animals. Matteson (1877, pp. 484-435) reports that the aplodontia
in Oregon ‘“‘is generally known as Mountain Boomer from his habit
of making a kind of booming noise.’? Lum (1878, pp. 10-13) has
never known it to make any sound by day or night, ‘‘save a kind of
growl when caught in a trap.”’ Price (1894, pp. 315-332) speaks of
a shrill cry heard several times when he was near a colony of mountain
beaver.
ENEMIES
Most writers agree with my own conclusion that aplodontia is a
very timid animal, but that it will fight fiercely when in a trap. Its
clumsy movements seldom permit of its inflicting the injury that its
strong jaws and teeth are well fitted to produce. When in pain, milky
white tears of a sticky nature cover the small eyes of the creature.
A pronounced lachrymal development in a burrowing animal must be
a useful feature.
Traps set in aplodontia burrows have caught weasels, skunks
(both Spilogale and Mephitis), mink, and gray fox (Dixon, MS,
1909) ; and coons (Lord, 1866, pp. 346-358), badgers, wildeats, fishers
(Lum, 1878, pp. 10-13), and owls are to be regarded as possible ene-
mies. Red squirrels, rabbits, and ground squirrels share the animals’
burrows or forage about their litter.
Cooper (1860, p. 82) says that the school children at Astoria,
Oregon, used to catch the awkward creatures by running them down.
This may indicate some activity in the daytime for this species. The
gait when running is a sort of gallop like that of a bear (Storer, MS,
1915). Observations upon an aplodontia in captivity have shown that
the creature runs as easily backwards as forwards; and this faculty
must be useful in the narrow burrows.
Economic IMpoRTANCE AND TRAPPING
In a few localities aplodontia does some damage by eating crops
and interfering with cultivation at the edges of the fields. Grinnell
(MS, 1912) records it as causing trouble along the line of the Union
Pacific Railroad over the central Sierra Nevada ‘‘by burrowing in the
diteh walls.’’ Shelton (MS, 1916) says that, in western Oregon,
534 University of California Publications in Zoology | Vou. 17
aplodontia makes itself a nuisance to foresters by undermining trails
and causing washouts.
Mountain beaver are easily captured and the usual method of trap-
ping them is to set No. 0 or No. 1 steel traps in the tunnel entrances.
The Indians (in Oregon and Washington), says Cooper (1860, p. 82),
catch the animals with ‘‘stone fall traps.’’ He adds that the skins
were not then being bought by the Hudson’s Bay Company. Gibbs
(in Suckley, 1860, pp. 100-106) relates how the Indians, probably the
Nisqually tribe in Washington, ‘‘trap and eat them and make gar-
ments by sewing the dried skins together.’’ The animals were there
resembling the figure four trap.
ce Le)
caught in traps Skins of the
California species bring only from eight to ten cents in the market, so
there is little danger that the mountain beaver will be exterminated
as a result of the fur trade.
Aplodontia is not hardy and if injured in the least does not live
long in captivity.
Transmitted February 4, 1918.
.
1918] Camp: Burrows of the Rodent Aplodontia 535
LITERATURE CITED
ANTHONY, H. E.
1916. Habits of Aplodontia. Bull. Amer. Mus. Nat. Hist., 35, 53-63, 8 figs.
AubDuBON, J. J., and BACHMAN, J.
1854. The quadrupeds of North America. (New York, Audubon), 3, vi+348,
BRETHERTON, B. J.
1895. Some Oregon mammals: The mountain boomer (Haplodon rufus).
Brooks, A.
1899. The sewellel, Aplodontia rufa. Recreation, 2, 258-259, 1 half tone.
Cooprr, J. G.
1860. Report upon the mammals collected on the Survey, U. S. Pae. R. R.
Expl. and Surv., 12, Zool. Rep., bk. 2, pt. 3, no. 2, chap. 1, pp. 73-88.
Cougs, E.
1877. ‘‘Haplodontidae’’ in Monographs of North American Rodentia, in
Rep. U. 8. Geol. Surv. Terr., 11, no. 9, pp. 543-601, 1 pl.
Evuiot, D. G.
1899. Catalogue of mammals from the Olympic Mountains, Washington,
with descriptions of new species. Field Columb. Mus. Zool., 1,
241-276, pls. 41-61, several unnumbered figs. in text.
1903. A list of mammals obtained by Edmund Heller, collector for the
Museum, from the coast region of northern California and Oregon.
Field Columb. Mus. Zool., 3, 175-198.
KELLOGG, L.
1916. Report upon mammals and birds found in portions of Trinity, Siskiyou
and Shasta counties, California. Univ. Calif. Publ. Zool., 12,
335-398, pls. 15-18, 1 fig. in text.
Lewis, M., and Cuark, W.
1814. History of the Lewis and Clark Expedition. (Paul Allen edition,
Philadelphia, Bradford), 2, ix + 522.
1904-5. Original journals of the Lewis and Clark Expedition, 1804-1806.
8 vols. (Thwaites’ edition, New York, Dodd), 8 [Atlas], i-xvi,
54 maps.
Lorp, J. K.
1866. The naturalist in Vancouver Island and British Columbia. 2 vols.
(London, Bentley), 1, xiv + 358, frontisp., 6 illus. in text.
Lum, S. K.
1878. The sewellel or show’tl. Amer. Nat., 12, January, 10-13.
Lyon, M. W., Jr.
1907. Notes on mammals collected at Mount Rainier, Washington. Smith-
sonian Mise. Coll., 50, 89-92.
Marteson, F. 8.
1877. The mountain boomer, or showtl. Amer. Nat., 11, 434-435.
Merriam, ©. H.
1886. Description of a new species of Aplodontia (Aplodontia major, sp.
nov.) from California. Ann. N. Y. Acad. Sci., 3, no. 10, pp. 312-
328, 2 pls., 1 fig. in text.
1899. Results of a biological survey of Mount Shasta, California. U.S.
Dept. Agr., Div. Biol. Surv., N. Amer. Fauna, 16, 1-179, pls. 1-5,
46 figs. in text.
536 University of California Publications in Zoology [ Vou. 17
Mur, J.
1909. Our national parks. (Boston, Houghton), x + 382 pp., numerous un-
numbered plates.
MourpaHy, J. M.
1876, The hunting fields of the Pacific Coast. Capturing the showtl. Rod
and Gun, May 20, p. 121.
PEALE, T. R.
1848. Mammalia and ornithology. U. S. Expl. Exped. (Wilkes) (Phila-
delphia, Sherman), 8, i-xxvi, 17-338, several unnumbered figs. in
, text.
PRICE, W. W.
1894. Notes on a collection of mammals from the Sierra Nevada Mountains.
Zoe, 4, 315-332.
RICHARDSON, J.
1829. On Aplodontia, a new genus of the order Rodentia, constituted for
the reception of the sewellel, a burrowing animal which inhabits
the northwestern coast of America. Zool. Jour., 4, 333-337.
STEPHENS, F.
1906. California mammals. (San Diego, West Coast Publishing Co.), 351
pp., frontisp., many unnumbered figs. in text.
SucKLEY, G.
1860. Report, in Zool. Rep., U. S. Pac. R. R. Expl. and Surv., 12, bk. 2, pt.
3, no. 2, chap. 2, pp. 89-106.
SuckLey, G., and Grpss, G.
1860. Op. cit., chap. 3, pp. 107-139, pls. 2, 5, 7, 9, 15.
TAYLOR, W. P.
1918. Revision of the rodent genus Aplodontia. Univ. Calif. Publ. Zool., 17,
435-504, pls. 25-29, 16 figs. in text.
TOWNSEND, C. H.
1887. Field-notes on the mammals, birds and reptiles of northern California.
Proce. U. 8. Nat. Mus., 10, 159-241.
INDEX*
Titles of papers in this volume, and names of new species, are printed in a
bold-faced type
Alaska, notes on some bats of, 431.
Allen, J. A., 226
Allen, Glover M., 226.
Ambystoma ensatum, 139; map show-
ing distribution in California,
131.
macrodactylum, 138; map showing
distribution in California, 131.
paroticum, 139; map showing distri-
bution in California, 131.
tigrinum, 138; map showing distri-
bution in California, 131.
Ammospermophilus nelsoni amplus,
15, 16.
Ammospermophilus nelsoni nelsoni,
15, 16; breeding habits, 19;
crowded out by Citellus beecheyi,
20; distribution of, 17; eco-
nomic relations, 19; effect of
farming on the status of, 19; life
history of, 17; map showing
range of, 18.
Ammospermophilus nelsoni nelsoni
Merriam, Notes on; with an Ac-
count of a New Spermophile
from the San Joaquin Valley,
California, 15.
Amphibians and Reptiles of Califor-
nia, Distributional List of, 127.
Amphibians, index to names of Cali-
fornia species, including all syn-
onyms, 201-208.
Aneides ferreus, 135; map showing
range in California, 133.
iécanus, 135; map showing distribu-
tion in California, 133.
lugubris farallonensis, 135; map
showing distribution in Califor-
nia, 133.
lugubris lugubris, 134; map show-
ing distribution in California,
133.
Anniella pulchra nigra, 170.
pulehra pulehra, 170.
Anser albifrons, Study of the Races
of the White-fronted Goose Oc-
curring in California, 209.
Anser albifrons, 209; measurements
of the races of, 220; occurring in
California, 209.
albifrons, 213, 215, 216, 220,
ventral view of, opp. 222.
gambeli, 213, 214, 215, 216,
220, ventral view of, opp. 222.
erythropus, 211.
Antrozous, 347.
* Univ. Calif. Publ. Zool., vol. 17.
pacificus, 352-357, 249; dorsal view,
opp. 390; dorsal view of skull of,
opp. 402; front view of head,
347; side view of skull of, opp.
404; map showing distribution,
351.
pallidus, 247, 249, 348-352; dorsal
view of skull of, opp. 402; side
view of skull of, opp. 404; map
showing distribution, 351.
Aphelocoma, compared with Siebero-
citta, 407; map showing distribu-
tion on Pacific Coast, 411; meas-
urements of, opp. 422.
Aphelocoma californica, 406, 407, 409,
421; variable features in, opp.
408.
californica, 410, 411, 412, 416.
immanis, 408, 411, 415, 416.
oocleptica, 411, 412, 413.
cyanea, 406.
hypoleuca, 406, 408, 411, 420; vari-
able features in, opp. 408.
insularis, 411, 418.
sieberi, 406.
sumichrasti, 407.
texana, 407.
woodhousei, 406, 411, 417.
unicolor, 407.
Aphelocoma, the Pacific Coast Jays
of the Genus, 405.
Aplodontia, age variation in, 439;
breeding habits, 532; burrows of,
519; drainage of, 531; economic
importance of, 533; enemies of,
533; feeding habits of, 528;
food of, 525; geographic distri-
bution of the nine recognized
races of, 455; geographic varia-
tion of, 445; habitat of, 519;
habits of, 450; hibernation of,
530; individual variation of,
441; list of species and sub-
species with type localities of,
453; molt and seasonal variation
in, 445; present systematic status
of, 450; sanitation of, 531; scat-
ology of, 531; sense perception
of, 532; sexual differences in,
440; trapping of, 533; variations
in the generic name of, 452.
Aplodontia, Brown, 454.
Humboldt, 470.
Mount Rainier, 465.
Northern, 463.
Olympic, 460.
[ 537 J
Index
Pacifie, 467.
Point Arena, 479.
Point Reyes, 480.
Sierra, 473.
Aplodontia californica, 531.
columbiana, 437.
chryseola, 437.
humboldtiana, 437.
leporina, 437
major, 437.
rainieri, 437.
nigra, 437.
olympica, 437.
pacifica, 437.
phaea, 437.
rufa californica, 522, 525; burrow
system of, 523; habitat of,
524; life study of, 531; map
showing distribution, 455.
columbiana, 522, 464; map show-
ing distribution, 455,
grisea, 437.
humboldtiana, 521, 522; map
showing distribution, 455.
nigra, 479, 521; map showing dis:
tribution, 455.
olympica, 460; map showing dis-
tribution of, 455.
pacifica, 467; map showing dis-
tribution of, 455.
phaea, 480, 531; map showing
distribution of, 455; nest-
galleries of, 521; plan of
underground tunnels, 520.
rainieri, 522, 465; map showing
distribution of, 455.
rufa, 437, 454, 522, 525; map
showing distribution of, 455.
Aplodontia, Excavations of Burrows
of the Rodent, with Observa-
tions on the Habits of the Ani-
mal, 517.
Aplodontia, Revision of the Rodent
Genus, 435.
Aplodontiidae, history of the, 447.
Arizona elegans, 192.
Ascaphus truei, 139.
Bailey, Vernon, 438.
Bangs, Outram, 438.
Bat, big-eared, little, 292.
Brown, Alaska, 267.
La Grulla, 290.
Large, 314; dorsal view of, opp.
392.
California, origin of, 240.
Little, 279; dorsal view of, opp.
392.
cave, 259.
foliage, oak, 285.
free-tailed, Mexican, 361; cluster
of, opp. 384.
Tacubaya, 357.
fringed, 297.
hoary, 330; dorsal view of, opp.
390; ventral view of, opp. 388.
Hollister, 261.
leaf-nosed, California, 252; cave of,
opp. 384.
long-legged, interior, 271.
northwestern, 267.
lump-nosed, dorsal view of, opp.
390.
intermediate, 344.
pale, 340.
mastiff, dorsal view of, opp. 394;
head of, opp. 396; ventral view
of, opp. 394.
California, 370.
Merriam, 311; dorsal view of, opp.
392.
Miller, 278.
pallid, desert, 348.
little, 288.
Pacific, 352; dorsal view of, opp.
390.
pocketed, 360.
red, dorsal view of, opp. 392.
western, 323.
Sierra, high, 9, 263; dorsal view of,
opp. 392.
silvery-haired, 300.
spotted, 336; dorsal view of, opp.
388.
Tejon, 276.
western, 305.
Yuma, 273.
Bats, 223; age variation of, 244;
classification of, 245; conserva-
tion of, 238; dentition of, 243;
economic value of, 238; enemies
of, 237; key to California, 246;
geographic distribution of, 241;
habits of, 232-236; measure-
ments of, 227; average measure-
ments of the California species
of, 251; modes of preserving,
228; nomenclature of, 245; senses
of, 228-232; sexual variation of,
244; voice of, 236.
Bat, A New Species of the Genus
Myotis from the High Sierra
Nevada of California, 9.
Bats, Notes on Some, from Alaska
and British Columbia, 431.
Bats, A Synopsis of the Species of
[ 538 ]
California, 223.
Batrachoseps attenuatus, 136; map
showing range of, 137.
major, 136; map showing range
of, 137.
pacificus, 136; map showing range
of, 137.
Beavers, 75; astragalus of, 95; bones
of the limbs and of the pectoral
and pelvie girdles, tabulation of
the differences noted in, 96-98;
caleaneum of, 95; clavicle of, 89;
femur of, 92; fibula of, 94; hum-
erus of, 89; os innominatum of,
92; radius of, 91; ribs of, 88;
scapula of, 88; sternum of, 88;
tibia of, 93; ulna of, 91; verte-
Index
brae: caudal of, 87; cervical of,
78; lumbar of, 80; sacral of, 86;
thoracic of, 80. SEE ALSO Castor.
Beavers, Osteological Relationships
of Three Species of, 75.
Boa, California, 177.
Borell, Adrey, 226.
Borell, Raleigh, 226.
Bovard, J. F., 438.
Branta canadensis, 211.
British Columbia, notes on some bats
of, 431.
Bryant, H. C., 21, 209.
Bufo alvarius, 144; map showing
range of, 141.
boreas, 61, 115.
boreas, 60, 115, 116, 143; map
showing range of, 141.
halophilus, 60, 61, 142; habitat
of, 62; map showing range
of, 141.
nelsoni, 115, 117, 143; map show-
ing range of, 141.
canorus, 59, 60, 61, 143; drawing
of, 60; habitat of, 62; map
showing range of, 141.
cognatus californicus, 141; map
showing range of, 141.
cognatus, 140; map showing
range of, 141.
columbiensis, 116.
halophilus, 115.
punctatus, 144; map showing range
of, 141.
woodhousii, 142; map showing
range of, 141.
Bufo canorus, A New Toad from the
Yosemite National Park, De-
scription of, 59.
Bush-tit, 419.
Camp, C. L., 2, 11, 59, 63, 115, 127,
226, 517.
Callisaurus ventralis myurus, 70.
ventralis, 70, 151.
Callospermophilus chrysodeirus per-
pallidus, 429.
Castor canadensis, 96.
belugae, 76, 81, 84,85; clavicle of,
opp. 106; femur of, opp. 114;
fibula of, opp. 106; humerus
of, opp. 108; os innominatum,
opp. 112; radius of, opp. 110;
sacral vertebrae of, opp.
102; seapula of, opp. 104;
thoracic vertebrae of, opp.
100; ulna of, opp. 110.
frondator, 77.
leucodonta, 76, 77, 79; clavicle
of, opp. 106; femur of, opp.
114; fibula of, opp. 106;
humerus of, opp. 108; radius
of, opp. 110; sacral verte-
brae of, opp. 102; scapula of,
opp. 104; thoracic vertebrae
of, opp. 100; ulna of, opp.
110.
phaeus, 76, 78; clavicle of, opp.
106; femur of, opp. 114;
fibula of, opp. 106; humerus
of, opp. 108; radius of, opp.
110; sacral vertebrae of,
opp. 102; scapula of, opp.
104; thoracie vertebrae of,
opp. 100; ulna of, opp. 110.
fiber, 77, 78, 81, 84, 85, 96; clavicle
of, opp. 106; femur of, opp.
114; fibula of, opp. 106; hu-
merus of, opp. 108; os innom-
inatum of, opp. 112; radius of,
opp. 110; sacral vertebrae of,
opp. 102; seapula of, opp. 104;
thoracic vertebrae of, opp.
100; ulna of, opp. 110.
subauratus, 77, 78, 79, 81, 84, 85,
96; clavicle of, opp. 106; fe-
mur of, opp. 114; fibula of,
opp. 106; humerus of, opp. 112;
os innominatum of, opp. 112;
radius of, opp. 110; sacral
vertebrae of, opp. 102; scapula
of, opp. 104; thoracie verte-
brae of, opp. 100; ulna of, opp.
110.
Charina bottae, 177; map showing
range in California, 178.
Chickadee, mountain, map showing
distribution in. California, 506;
subspecies of, 505.
Bailey, 511.
Inyo, 509.
Rocky, 508.
short-tailed, 510.
Chilomeniseus cinetus, 182; map
showing range in California, 185.
Chipmunk, Kern Basin, 5.
Los Banos Antelope, 15.
Mariposa, 4.
Mono, 3.
Chiroptera, 252.
Chuckwalla, 153.
Citellus beecheyi, 20.
Clemmys marmorata, 199.
Cnemidophorus hyperythrus beldingi,
175; map showing range in Cali-
fornia, 173.
stejnegeri, 71.
tigris, 68; subspecies of, 71; varia-
tion in, 71.
mundus, 71, 173; map showing
range in California, 173.
stejnegeri, 174; map showing
range in California, 173.
tigris, 172; map showing range in
California, 173.
undulatus, 71.
Coleonyx variegatus, 149; map show-
ing range in California, 167.
Coluber constrictor vetustus, 189;
map showing range in California,
191.
flagellum frenatus, 190; map show-
ing range in California, 191.
[ 539 J
Index
lateralis, 190; map showing range
in California, 191.
taeniatus, 192; map showing range
in California, 191.
Contia mitis, 183; map showing range
in California, 185.
Cony, Yosemite, 6.
White Mountains, 429.
Corynorhinus, 339.
rafinesquii intermedius, 247, 249,
344-347; dorsal view of skull
of, opp. 402; map showing dis-
tribution of, 342; side view of
skull, opp. 404.
pallescens, 247, 249, 340-344;
asleep, photograph of, opp.
390; map showing distribu-
tion of, 342.
Crotalus atrox, 196; map showing
range in California, 195.
cerastes, 198; map showing range
in California, 195.
exsul, 196; map showing range in
California, 195.
mitchellii, 197; map showing range
in California, 195.
oreganus, 194; map showing range
in California, 195.
tigris, 197; map showing range in
California, 195.
Crotaphytus baileyi, variation in, 70.
collaris baileyi, 70, 153.
silus, 152.
wislizenii, 152; variation in, 70.
Dermochelys schlegelii, 198.
Description of Bufo canorus, A New
Toad from the Yosemite Na-
tional Park, 59.
Diadophis amabilis, 184; map show-
ing range in California, 185.
Diagnosis of Seven New Mammals
from East-Central California, 1;
literature cited, 8.
Dickey, D. R., 257.
Dipso-saurus dorsalis, 150.
Distributional List of the Amphi-
bians and Reptiles of California,
127.
Eptesicus, 314.
fuscus, 247, 249, 314-322; collecting
site of, opp. 384; dorsal view
of, opp. 392; dorsal view of
skull of, opp. 402; map show-
ing occurrence of in California,
319; side view of skull of, opp.
404; wing of, 246.
Euderma, 336.
maculatum, 247, 249, 336-339; map
showing distribution of, 303;
photograph from dried skin,
opp. 338.
Eumeces gilberti, 72.
quinquelineatus, 73.
skiltonianus, 68, variation in, 72.
Eumops, 369.
californicus, 246, 249, 370-376;
dorsal view of, opp. 394; dorsal
view of skull of, opp. 396; head
of, opp. 396; side view of
skull of, opp. 396; ventral
view of, opp. 394.
Eutamias amoenus amoeuus, 3, 4.
monoensis, 3.
merriami kernensis, 5, 6.
mariposae, 4.
merriami, 4, 6.
Evermann, B. W., 209, 226.
Excavations of Burrows of the Rod-
ent Aplodontia, with Observa-
tions on the Habits of the Ani-
mal, 517; literature cited, 535.
Frogs, key to California, 123; map
showing distribution in Califor-
nia, 147.
Frogs of California, Notes on the
Systematic Status of the, 115.
Frog, leopard, 149.
red-legged, California, 148.
Oregon, 148.
spotted, Nevada, 148.
western, 148.
tree, Arizona, 145.
Pacific, 144.
yellow-legged, California, 117, 146.
Sierra Madre, 118, 147.
Sierra Nevada, 120, 146.
western, distribution of, 124.
Gecko, map showing distribution in
California, 167.
banded, 149.
Gerrhonotus coeruleus, 169; map
showing range in California,
167.
palmeri, 168; map showing range
in California, 167.
scincicauda, 68.
scincicauda, 166; map showing
range in California, 167.
webbii, 168; map showing range
in California, 167.
Gidley, J. W., 438.
Gilbert, C. H., 225.
Goose, White-Fronted (Anser albi-
frons), A Study of the Races
Occurring in California, 209.
Gopher, pocket, Mohave River, 427.
White Mountains, 425.
Grinnell, H. W., 9, 223, 431.
Grinnell, Joseph, 1, 15, 23, 59, 127,
226, 423, 438, 505, 518.
Habits and Food of the Roadrunner
in California, 21; literature
cited, 48.
Hanna, W. C., 226.
Henshaw, Judge F. W., 209, 210, 218.
Henshaw, H. W., 226.
Henshaw, Samuel, 438.
Holden, F. H., 75.
Hollister, Ned, 226.
Howell, A. H., 226.
[ 540 ]
Index
Hyla arenicolor, 145.
regilla, 144.
Hypsiglena ochrorhynchus, 188; map
showing range in California, 178.
Iguana, desert, 150.
Index to names of California am-
phibians and reptiles, including
all synonyms, 201—208.
Jackson, Hartley H. T., 438.
Jays of the Pacific Coast
Genus Aphelocoma, 405.
Kinosternon sonoriense, 200.
Kofoid, C. A., 226.
Lampropeltis boylii, 186.
ealiforniae, 187.
conjuneta, 187.
pyromelana multicincta, 184.
Lasionycteris, 299.
noctivagans, 247, 249, 300-304;
collecting site of, opp. 384;
dorsal view of skull of, opp.
398; map showing distribution,
303; side view of skull of, opp.
400.
Leptotyphlops humilis, 176.
Lichanura roseofusea, 177; map show-
ing range in California, 178.
Literature cited, 8, 14, 20, 48, 74,
99, 125, 221, 422, 433, 376-382,
484, 515, 535.
Lizard. alligator, map showing dis-
tribution in California, 167.
California, 166.
Coast, 169.
San Diego. 168.
Sierran, 168.
blue-bellied, map showing distri-
bution in California, 161.
island, 162
Tenaya, 66, 160.
brown-shouldered, desert, 155.
northern, 154.
southern, 156.
collared, Bailey, 153.
fence, 160.
footless, black, 170.
silvery, 170.
gridiron-tailed, southern, 151.
leopard, Great Basin, 152.
San Joaquin, 152.
Mearns, 154.
mountain, 157.
night, desert, 171.
Henshaw, 171.
island, 172.
orange-throated, Belding, 175.
rough-sealed, desert, 162.
dusky, 163.
sand, ocellated, 150.
Van Denburgh, 67, 159.
whip-tailed, map showing distri-
bution in California, 173.
California, 173.
desert, 172.
Stejneger, 174.
Lizards, California, an Account of
of the
[ 541 J
the Subspecies of Sceloporus oc-
cidentalis with a Description of
a New Form from the Sierra
Nevada and Systematic Notes on
Other Forms, 63.
Macrotus, 252.
ealifornicus, 246, 249,
cave of, opp. 384; dorsal view
of skull of, opp. 402; map
showing distribution, 256; side
view of skull of, opp. 404.
Mailliard, J. W., 405, 505.
Mailliard, Joseph, 405, 505.
Mammals, Diagnoses of Seven New,
from East-Central California, 1.
Mammals, Six New Species from the
Mohave and Inyo Regions of
California, 423.
Marten, Sierra Pine, 2.
Martes caurina caurina
sierrae, 2.
Merriam, C. H., 438.
Merriam, J. C., 437.
Microchiroptera, 252.
Miller, G. S., 226, 438.
Mohave Desert, six new
from, 423.
Mole, Mono, 423.
San Joaquin, 1.
Molossidae, 357.
Moreom, G. F., 505.
Muir, John, 7.
Myotis, 258.
ealifornicus ecalifornicus, 248, 250,
279-285; collecting site of,
opp. 384; dorsal view of, opp.
392; dorsal view of skull of,
opp. 398; map showing dist-
tribution of, 284; side view of
skull of, opp. 400.
eaurinus, 432.
pallidus, 248, 250, 288-290; dor-
sal view of skull of, opp.
398; map showing distribu-
tion, 284; side view of skull
of, opp. 400.
quereinus, 248, 250, 285-288; map
showing distribution in Cali-
fornia, 284.
evotis, 247, 250, 292-297; dorsal
view of skull of, opp. 398; map
showing distribution in Cali-
fornia, 296; side view of skull
of, opp. 400.
longierus interior, 248, 250, 271—
273; map showing distribu-
tion in California, 270.
longicrus, 248, 250, 267-271, 431;
collecting site of, opp. 384;
dorsal view of skull of, opp.
398; map showing distribu-
tion in California, 270; side
view of skull of, opp. 400.
lucifugus alascensis, 248, 250, 267,
432; map showing distribution
in California, 256.
252-258;
9
) ae
mammals
25
Index
altipetens, 248, 250, 263-267;
collecting site of, opp. 384;
dorsal view of, opp, 392;
dorsal view of skull of, opp.
398; map showing distribu-
tion in California, 256; side
view of skull of, opp. 400.
oceultus, 248, 249, 250, 261-263;
dorsal view of skull of, opp.
398; map showing distribution
of, 303; side view of skull of,
opp. 400.
orinomus, 248, 250, 290-292; dorsal
view of skull of, opp. 398;
map showing distribution in
California, 303; side view of
skull of, opp. 400.
thysanodes, 248, 250, 297-299; col-
lecting site of, opp. 3°84; dorsal
view of skull of, opp. 398; map
showing distribution in Cali-
fornia, 296; side view of skull
of, opp. 400.
yumanensis altipetens, 9.
saturatus, 248, 250, 278-279; map
showing distribution in Cali-
fornia, 275.
sociabilis, 9, 248, 250, 276-278;
dorsal view of skull of, opp.
398; map showing distribu-
tion in California, 275; side
view of skull of, opp. 400.
yumanensis, 248, 250, 273-276;
map showing distribution in
California, 275.
velifer, 248. 259-261; dorsal view
of skull of, opp. 398; map
showing distribution in Cali-
fornia, 296; side view of skull
of, opp. 400.
Neale, George, 209.
Nelson, Edward W., 226, 438, 505.
New Bat of the Genus Myotis from
the High Sierra Nevada of Cali-
fornia, 9.
Newt, Pacific Coast, 130.
Notes on Some Bats from Alaska and
British Columbia, 431; literature
cited, 433.
Notes on the Systematic Status of
the Toads and Frogs of Califor-
nia, 115; literature cited, 125.
Notophthalmus torosus, 130; map
showing distribution in Califor-
Nia loile
Nycteris, 322.
borealis teliotis, 247, 249, 323-330;
dorsal view of, opp. 392; dorsal
view of skull of, opp. 402;
female with three young, opp.
392; map showing distribution
in California, 327; side view
of skull of, opp. 404.
einerea, 247, 249, 330-336; dorsal
view of, opp. 390; dorsal view
of skull of, opp. 402; front
view of, opp. 388; map show-
ing distribution in California,
327; side view of skull of, opp.
404; specimen secured at Fresno,
opp. 388.
Nyctinomus, 357.
depressus, 247, 249, 357-360.
femorosaccus, 246, 249, 360-361;
map showing distribution in
California, 303.
mexicanus, 247, 249, 361-369; eolony
of, opp. 384; dorsal view of
skull of, opp. 402; map show-
ing distribution in California,
366; side view of head of, 362;
side view of skull of, opp. 404.
Nyctophilinae, 347.
Ochotona schisticeps albatus, 6.
muiri, 6.
sheltoni, 429.
schisticeps, 6.
Osgood, W. H., 438.
Osteological Relationships of Three
Species of Beavers, 75; literature
cited, 99.
Pacific Coast Jays of the Genus
Aphelocoma, 405; literature cited,
422.
Palmer, T. S., 438.
Penthestes atricapillus, 507.
gambeli, 505, 507; measurements of
the four races of, 513.
abbreviatus, 510, 512, 513; map
showing distribution of, 506;
measurements of, 514.
baileyae, 511, 512, 513; map show-
ing distribution of, 506;
measurements of, 514.
gambeli, 508, 512; measurements
of, 513.
inyoensis, 509, 512, 513; map
showing distribution of, 506;
measurements of, 514.
Perodipus mohavensis, 428.
Phrynosoma blainvillii, 120.
blainvillii, 164.
frontale, 165.
douglassii douglassii, 164.
m’eallii, 166.
platyrhinos, 165.
Phyllostomidae, 252.
Pipilo maculatus, 408, 419.
Pipistrellus, 304.
hesperus hesperus, 247, 249, 305-
311; dorsal view of skull of,
opp. 398; map showing dis-
tribution, 309; side view of
skull of, opp. 400.
merriami, 247, 249, 311-314; col-
lecting site of, opp. 384;
dorsal view, opp. 392; map
showing distribution of,
309.
Pituophis catenifer catenifer, 193.
deserticola, 193.
Plestiodon skiltonianum, 175.
[ 542 ]
Index
Plethodon croceater, 132; map show-
ing range in California, 133.
elongatus, 134; map showing range
in California, 133.
eschscholtzii, 132; map showing
range in California, 133.
Preble, E. A., 438.
Psaltriparus minimus, 419.
ealifornicus, 408.
plumbeus, 408.
Rabbit, Brush, Mariposa, 7.
Racer, map showing distribution in
California, 191.
red, 190.
striped, California, 190.
Nevada, 192.
yellow-bellied, western, 189.
Rana aurora, 115, 120.
aurora, 123, 124, 148; map show-
ing range in California, 147.
draytonii, 123, 124, 148; map
showing distribution in Cali-
fornia, 147.
boylii boylii, 117, 118, 119, 120,
122, 125, 146; map showing
range in California. 147.
muscosa, 118, 123, 147; map
showing range in Califor-
nia, 147.
sierrae, 120, 122, 123, 146; map
showing range in California,
147.
draytonii, 115.
pachyderma, 117.
pipiens, 149; map showing distri-
bution in California, 147.
pretiosa, 115, 117, 120.
luteiventris, 123, 124, 148; map
showing distribution in Cali-
fornia, 147.
pretiosa, 122, 123, 124, 148; map
showing distribution in Cali-
fornia, 147.
temporaria pretiosa, 117.
Rat, kangaroo, Mohave, 428.
Rattlesnake, map showing distribu-
tion in California, 195.
red, 196.
Pacifie, 194.
nallid, 197.
Texas, 196.
tiger, 197.
Reptiles and Amphibians of Califor-
nia, A Distributional List of,
127.
Reptiles, index of Californian, 201—
208.
Revision of the Rodent Genus Aplo-
dontia, 435; literature cited,
484-495,
Rhinocheilus lecontei, 188; map show-
ing range in California, 178.
Richmond, C. W., 505.
Roadrunner, 21; ¢arrying a whip-
tailed lizard, opp. 56; economic
status of, 43; eggs of, 27; food
of, 29; life history of, 24; nest
of, 26, opp. 52; nest in cactus,
opp. 54; range of, 25; stomach
capacity, of, 43; stomach con-
tents of, opp. 58; young of, 27.
Roadrunner, Habits and Food of, in
California, 21.
Rowley, John, 226.
Salamander, a New Alpine, from the
Yosemite National Park, Cali-
fornia, 11.
Salamander, Arboreal, 134.
British Columbia, 139.
Del Norte, 134.
Farallon, 135.
garden, 136.
island, 136.
long-toed, 138.
marbled, 139.
Mount Lyell, 11, 13, 132.
Oregon, 132.
rusty, 135.
Shasta, 135.
slender, 136; map showing distri-
bution in California, 137.
tiger, 138.
yellow-spotted, 132.
Salamanders, map showing distribu-
tion in California, 131, 133.
Salvadora hexalepis, 189; map show-
ing range in California, 178.
San Joaquin Valley, antelope chip-
munks of, 15.
Sauromalus ater, 153.
Scapanus latimanus campi, 1.
monoensis, 423; measurements
of, 424.
sericatus, 2.
Seaphiopus hammondii hammondii,
Sceloporus graciosus, 67; subspecies
of, 67; variation in, 67.
graciosus, 68, 157; map showing
range in California, 158.
vandenburgianus, 67, 159; map
showing range in California,
158. :
magister, 162; map showing range
in California, 158.
occidentalis, 68; subspecies of, 63;
variation in, 63.
becki, 162; map showing range
in California, 161.
bi-seriatus, 65, 66, 160; map
showing range in California,
161.
occidentalis, 65; map showing
range in California, 161.
taylori, 65, 66, 160; map show-
ing range in California, 161.
orcutti. 163; map showing range in
California, 158.
Shelton, A. C., 438, 518.
Sidewinder, 198.
Skiff, F. J. V., 438.
Skink, western, 175.
[543 ]
Index
Snake, map showing distribution in
California, 185.
burrowing, banded, 182.
desert, 182.
faded, 192.
garter, California, 181.
elegant, 180.
Marcy, 179.
Pacific Coast, 179.
western, 181.
gopher, desert, 193.
western, 193.
ground, Texas, 183.
king, black, 187.
Boyle, 186.
California, 187.
eoral, 184.
long-nosed, 188.
night, spotted, 188.
patch-nosed, 189.
ring-necked, western, 184.
rubber, 177.
sharp-tailed, 185.
worm, 176.
Snake. See also Rattlesnake, Side-
winder.
Sonora episcopa, 183; map showing
range in California, 185.
occipitalis, 182; map showing range
in California, 185.
Snyder, J. O., 225.
Spadefoot, western, 140.
Spelerpes leprosus, 12.
platycephalus, 11, 12, 132; drawings
of, 13; map showing range in
California, 133.
Spelerpes platycephalus, A New Al-
pine Salamander from the Yo-
semite National Park, Califor-
nia, 11; literature cited, 14.
Spermophile, A New Species from
the San Joaquin Valley, Califor-
nia, with Notes on Ammosper-
mophilus nelsoni nelsoni Mer-
riam, 15; literature cited, 20.
Squirrel, ground, Inyo golden-man-
tled, 429.
Stejneger, Leonhard, 130, 226.
Stone, Witmer, 226.
Storer, Loe 30:
Study of the Races of the White-
Fronted Goose (Anser albifrons)
Occurring in California, 209;
literature cited, 221.
Subspecies of the Mountain Chicka-
dee, 505; literature cited, 515.
Subspecies of Sceloporus occidentalis
with Description of a New Form
from the Sierra Nevada and
Systematic Notes on Other Cali-
fornia Lizards, 63; literature
eited, 74.
Swarth, H. 8., 15, 209, 405, 505.
Swerdfeger, Grace, 226.
Sylvilagus bachmani bachmani, 7.
cinerascens, 7.
mariposae, 7.
Synopsis of the Bats of California,
223; literature cited, 376-382.
Tantilla eiseni, map showing range
in California, 185.
Taylor, W. P., 9, 15, 435, 518.
Taverner, P. A., 226.
Testudo agassizii, 199; map showing
range in California, 167.
Toad, Arroyo, 141.
Bell, American, 139.
California, 142.
Colorado River, 144.
Great Plains, 140.
horned, Blainville, 164.
California, 165.
desert, 165.
flat-tailed, 166.
pigmy, 164.
Nevada, 143.
Northwestern, 143.
Rocky Mountain, 142.
spotted, 144.
Yosemite Park, 59, 143.
Toad, A New Form, Bufo canorus,
from the Yosemite National
Park, Description of, 59.
Toads, map showing distribution in
California, 141; key to Califor-
nia, 116.
Toads and Frogs of California, Notes
on the Systematic Status of, 115.
Thamnophis marcianus, 179.
ordinoides elegans, 180.
hammondii, 181.
ordinoides, 179.
sirtalis parietalis, 181.
Thayer, J. E., 405.
Thomomys perpallidus mohavensis,
427.
melanotis, 425; measurements of,
426.
Thryomanes bewicki, 408, 419.
Tortoise, desert, 199; map showing
distribution in California, 167.
Towhee, spotted, 419.
Turtle, leatherback, Pacific, 198.
mud, Arizona, 200.
Pacific, 199.
Uma notata, 150.
Uta graciosa, 156.
mearnsi, 154.
ornata, 157.
stansburiana, map showing distri-
bution in California, 155; sub-
species of, 68; variation in, 68.
elegans, 69, 155; map showing
distribution of, in Califor-
nia, 155.
hesperis, 69, 155; map showing
distribution in, California,
155.
stansburiana, 69, 154; map show-
ing distribution in Califor-
nia, 155.
[544]
Index
Uta, long-tailed, 156.
tree, 157.
Vespertilionidae, 258.
Vespertilioninae, 258.
Vireo, Hutton, 419.
Warren, E. R., 505.
Wear, Winifred N., 226.
White-Fronted Goose, A Study of the
Races Occurring in California,
209.
[ 545 J
Wren, Bewick, 419.
Xantusia henshawi, 171.
riversiana, 172.
vigilis, 171.
Yosemite, Natural History Survey
hay May alle bee
Yosemite National Park, Description
of a New Toad from, 59.
Yosemite National Park, A New Al-
pine Salamander, Spelerpes platy-
cephalus, from, 11.
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UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued)
62, 2 figures in text, November, 1915 2000 eager it)
5. Notes on the Tintinnoina. 1, On the Probable Origin of Dictyocysta tiara
Haeckel. 2. On Petalotricha entzi, sp. nov., by Charles Atwood Kofoid,
Pp. 63-69, 8 figures in text. December, 1915 00 — 05
6 BUA. ine Multiple Fission in: Hexamitus, by Olive Swezy. Pp. 71-88,
plates 9-11.
7. On a New ‘Trichomonad Flagellate, Trichomitus parvus, from the Intestine
of Amphibians, by Olive Swezy. Pp. 89-94, plate 12.
Nos. 6 and 7 in one cover. December,-1915 02 25
9. Three New Helices from California, by 8. Stillman Berry. Pp, 107-111.
10. On Trypanosoma triatomae, a New Flagellate from a Hemipteran Bug from
and Irene McCulloch. Pp. 113-126, piates 14-15. February, 1916 ...2..0.... 15
11. The Genera Monocercomonas and Polymastiz, by Olive Swezy. Pp. 127-138,
plates 16-17 -Pebrnaryy Ore Fo oe oa a ate |)
12. Notes on the Spiny Lobster (Panulirus interruptus) of the California Coast,
by Bennet M. Allen. Pp. 139-152, 2 figures in text, March, 1916 2... 15
18. Notes on the Marine Fishes of California, by Carl L. Hubbs. Pp. 153-169,
plates- 18-202 March. 1916 ee as SSSA 15
Nutrition by Organic Substances in Solution inthe Water, by Calvin O.
Esterly. Pp. 171-184, 2 figures in text. “March, 1916 2c tmage Wee |
15. The Kinetonucleus of Flagellates and the Binuclear Theory of Hartmann,
by Olive Swezy. Pp. 185-240, 58 figures in text, March, 1916-0... OO
16..On the Life-History of a Soil Amoeba, by Charlie Woodruff Wilson, Pp,
241-292= plates 18-25. Sony, 1916.20. See oe a, A ae hen 00
17. Distribution of Land Vertebrates of Southeastern Washington, by Lee
Raymond Dice. Pp, 293-348, plates 24-26. June; 1916 2.22 60
18. The Anatomy of Heptanchus maculatus: the. Endoskeleton, by J. Frank
Daniel. Pp, 349-370, pls. 27-29, 8 text figures. December, 1916 0.00222. 25
19. Some Phases of Spermatogenesis in the Mouse, by Harry B. Yocom. Pp.
S7F-880; -plates30;-°danuary,- 197 eso ee eh 10
20° Specificity in Behavior and the Relation between Habits in Nature and
Reactions in the Laboratory, by Calvin O. Esterly. Pp. 381-392, March,
21, The Occurrence of a Rhythm in the Geotropism of Two Species of Plank-
ton Copepods when Certain Recurring External Conditions are Absent, by
Calvin O. Esterly. Pp. 393-400. March; 1917 2o22..2.-jccscceci ccc ecedeeee te eeeetnne 10
22. On Some New Species of Aphroditidae from the Coast of California, by
Christine Essenberg. Pp. 401-480, plates 31-37, March, 1917 000. peek er)
23, Notes on the Natural History and Behavior of Emerita analoga (Stimpson),
by. Harold Tupper Mead. Pp. 431-438, 1 text figure, April, 1917 —......... ESS ALO:
24, Ascidians of the Littoral Zone of Southern California, by William E. Ritter
and Ruth A. Forsyth. Pp, 439-512, plates 38-46. August, 1917 -............. 1.00
2 Index in preparation.
Vol. 17, 1. Diagnoses of Seven New Mammals from East-Central California, by Joseph
Grinnell and Tracy I. Storer. Pp. 1-8. ;
2, A New Bat of the Genus Myotis from the High Sierra Nevada of Cali-
fornia, by Hilda Wood Grinnell. Pp, 9-10,
Nos. 1 and 2 in one cover, August, 1916 2....-20...2-c2ccce cee cecctn ee eee amiss |
$. Spelerpes platycephalus, a New Alpine Salamander from the Yosemite
National Park, California, by Charles Lewis Camp. Pp. 11-14.. Septem-
DF ata M6 5 Ret RA aN Aas abe aa LO EE gece Oe ee a ene 05
4, A New Snermophile from the San Joaquin Valley, California, with Notes
on Ammospermophilus nelsoni nelsoni Merriam, by Walter P. Taylor. Pp.
15-20, 1 figure in text. October, 1916.22.22... cle... ceerne . 05
6. Habits and Food of the Roadrunner in California, by Harold 0. Bryant.
Pp. 21-58, plates 1-4, 2 fignres in text. October, 1916 02. Meme.
6, Description of Bufo canorus, a New Toad from the Yosemite National Par: 3
by Charles Lewis Camp. Pp. 59-62, 4 figures in text. November, 1916... .05
7. The Subspecies of Sceloporus occidentalis, with Description of a New Form
from the Sierra Nevada and Systematic Notes on Other California
Lizards, by Charles Lewis Camp. Pp. 63-74. December, 1916... “10
UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued)
. Osteological Relationships of Three Species of Beavers, by F. Harvey
Holden. . Pp. 75-114, plates 6-12, 18 text figures. March, 1917 2...0.......2.
. Notes on the Systematic Status of the Toads-and-Frogs of California, by
Charles Lewis Camp. Pp. 115-125, $3 text figures. February, 1917 0.0...
10: A Distributional List of the Amphibians and Reptiles of California, by
Joseph Grinnell and Charles Lewis Camp. Pp. 127-208, 14 figures in text.
Pip Uh Bes 2) Bs false oo es ects ED a SR SoC etiam oh ate emery kee Me, Seen
11. A Study of the Races of the White-Fronted Goose (Anser albifrons) Oceur-
Ting in. California, by H. S. Swarth and Harold C. Bryant. Pp. 209-222,
2 ‘figures in text, plate 13.- .October, 1917 =. - oe ee
12. A Synopsis of the Bats of California, by Hilda Wood Grinnell. Pp. 223-404,
plates 14-24, 24 text-figures. January 31, 1918
13. The Pacific Coast Jays of the Genus Aphelocoma, by a. §.-Swarth. Pp.
405-422, 1 figure in text, February 23, 19V8 2202 cnc csccectecnsenceecnnaee
14, Six New Mammals from the Mohave Desert: and Tnyo Regions of California, ;
by Joseph Grinnell. . Pp. 423-430.
15. Notes on Some Bats from Alaska and British Columbia, by Bids Wood -
Grinnell. Pp. 431-433. :
Nos.-14 and-15 in one cover.: April, 1918 aa ee Arche Sir i a eee tee
16. Revision of the Rodent Genus Aplodontia, by Walter P. Taylor. Pp. 435- -
504, plates 25-29, 16 text figures. May, 1918) _.c2.u icici ected
17. The Subspecies of the Mountain Chickadee, by Joseph Grinnell.. Pp. 505-
515539 text: fisures. Mays 2918s nos SE ae i Soca peers
18. Excavations of Burrows of the Rodent Aplodentia, with Observations on
the Habits of the Animal, by Charles Lewis Camp. Pp. 517-536, 6 figures
intext dune, T0184 ko ee Oe ee Ree Sees see Sa
Vol 18. 1, Mitosis in Giardia microti, by William C. Boeck... Pp. 1-26, plate 1. Octo-
bey, QE 3S ee Se ra aS a es
2. An Unusual Extension of the Distribution of the Shipworni in San Fran-
cisco Bay, California, by Albert L. Barrows. Pp. 27-43. December, 1917.
8. Description of Some New Species of Polynoidae from the Coast of Cali-
fornia, by Christine Essenberg. Pp. 45-60, plates 2-3. October, 1917.
4. New Species of Amphinomidae from the Pacific Coast, by Christine Essen-
5
o ow
berg. Pp. 61-74, plates 4-5. October, TOU7 cen. ceec ieee cesnetteatdeceteceeeee ten
. Crithidia euryophthalmi, sp. nov., from the Hemipteran Bug, Euryophthalmus
convivus Stal, by Irene McCulloch. Pp. 75-88, 35- text figures. Decem-
DOES POET en a ane ae rea ESL ey creat
6. On the Orientation of Hrythropsts, by Charles Atwood Kofoid and Olive
Swezy. Pp. 89-102, 12 figures in text. Deceniber, 1917 ........ ae Ree ate
7. The Transmission of Nervous Impulses in Relation to Locomotion ‘in the 3
Earthworm, by John F. Bovard. Pp. 103-134, 14 figures in text. January, =
Bi PA: er on Ma eA Pea tH A can BN fe aN ey CE Ey EEE Se ayn ee Ee 3
8. The Function of the-Giant Fibers in Earthworms, by. John F. Bovard. Pp.
135-144, 1 figure in text. January, 1918 -.2...2 oo
9. A Rapid Method for the Detection: of Protozoan Cysts in “Mammalian
Faeces, by William C. Boeck. -Pp. 145-149. December, 1917 ..........
10. The Musculature of Heptanchus maculatus, by Pirie-Davidson... Pp, 151-170,
12 figures in-text. March, 1948 os eek cachet cca cto tke otadeoeres
11. The Factors Controlling the Distribution-of the Polynvidas of the Pacific
Coast of North America, by Christine Essenberg. Pp. alae plates 6-8,
2 figures-in “text. Merch,’ 19782-0528 ae See es CRs canada ugeen :
12. Differentials in Behavior of the Two Generations of Salea democratica
Relative to the Temperature of the Sea, by Ellis L. Michael. Pp. 239-298;
plates 9-11, 1 figure in text. March, 1918 2ncccloco.tccccen ccc ceeetiviepecenecneeennene cee
18. A Quantitative Analysis of the Molluscan Fauna of San Francisco Bay, by ©
EB. L. Packard. Pp. 299-336, a 12-13, 6 figs. in text. April, 1918...
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